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Full text of "A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands"

n Friedlander 



NOAA Center for Coastal Monitoring 

Biogeography Branch 



NOAA Office of National Marine Sanctuaries 
Papahanaumokuakea Marine National Monument 



NOAA Center for Coastal Monitoring and Assessment 

Biogeography Branch 

University of Hawaii at Manoa 



NOAA Center for Coastal Monitoring and Assessment 

Biogeography Branch 





NOAA Center for Coastal Monitoring and Assessment 

Biogeography Branch 




NOAA TECHNICAL 



R A N D U 



NOS NCCOS 




<S3 



Citations 

Citation for entire document: 

Friedlander, A., K. Keller, L. Wedding, A. Clarke, M. Monaco (eds.). 2009. A Marine Biogeographic Assess- 
ment of the Northwestern Hawaiian Islands. NOAA Technical Memorandum NOS NCCOS 84. Prepared by 
NCCOS's Biogeography Branch in cooperation with the Office of National Marine Sanctuaries Papahanau- 
mokuakea Marine National Monument. Silver Spring, MD. 363 pp. 

Example citation for an individual chapter (example of Oceanographic and Physical Setting chapter): 

Desch, A., T. Wynne, R. Brainard, A. Friedlander, J. Christensen. 2009. Oceanographic and Physical Setting, 
pp. 17-63. In: Friedlander, A., K. Keller, L. Wedding, A. Clarke, M. Monaco (eds.). 2009. A Marine Biogeo- 
graphic Assessment of the Northwestern Hawaiian Islands. NOAA Technical Memorandum NOS NCCOS 
84. Prepared by NCCOS's Biogeography Branch in cooperation with the Office of National Marine Sanctuar- 
ies Papahanaumokuakea Marine National Monument. Silver Spring, MD. 363 pp. 



Acknowledgements 

Project funding was provided by the Office of National Marine Sanctuaries Papahanaumokuakea Marine 
National Monument, the National Centers for Coastal Ocean Science and the Coral Reef Conservation 
Program. 

The completion of this document would not have been possible without the participation of the people rec- 
ognized below. Their efforts to compose, format, review and edit the document are very much appreciated. 
Particular recognition and thanks is extended to Jamison Higgins for her assistance with the layout and 
production of this publication. 

In addition to chapter authors, the editors would like to acknowledge the following contributors and 
reviewers: 

George "Bud" Antonelis, Jay Barlow, Jamison Gove, Francine Fiust, Sarah Hile, David Hyrenbach, John 
Klavitter, Kevin McMahon, Gustav Paulay, Audrey Rivero, Jenny Waddell, Susan White and Lee Ann Wood- 
ward. 



For more information 

For more information about this report, please contact CCMA's Biogeography Branch Chief Mark Monaco 
Ph.D, at (301) 713-3028 or visit http://ccma.nos.noaa.gov/about/biogeography/. 



Cover 

The covers were designed and created by Gini Kennedy (NOAA). Cover photos by James Watt (NOAA), 
James Maragos (U.S. Fish and Wildlife Service) and the Hawaii Volcano National Park. 



Mention of trade names or commercial products does not constitute endorsement or recommendation for 
their use by the United States government. 



A Marine Biogeographic Assessment of the 
Northwestern Hawaiian Islands 



Prepared for NOAA's Office of National Marine Sanctuaries (ONMS) 
Papahanaumokuakea Marine National Monument 



Biogeography Branch 

Center for Coastal Monitoring and Assessment (CCMA) 

National Centers for Coastal Ocean Science (NCCOS) 

NOAA's National Ocean Service 

1305 East West Highway (SSMC-IV, N/SCI-1) 

Silver Spring, MD 20910 

USA 



NOAA Technical Memorandum NOS NCCOS 84 



March 2009 



Editors 

Alan Friedlander, CCMA Biogeography Branch 

Kaylene Keller, ONMS Papahanaumokuakea Marine National Monument 

Lisa Wedding, CCMA Biogeography Branch; University of Hawaii at Manoa 

Alicia Clarke, CCMA Biogeography Branch 

Mark Monaco, CCMA Biogeography Branch 



4*0* 



K, 



National Oceanic and National Ocean Service 
Atmospheric Administration 

Mary M. Glackin John H. Dunnigan 

Deputy Under Secretary Assistant Administrator 



Table of Contents 



Executive Summary i 

Chapter 1: Introduction 1 

Alan Friedlander, Kaylene E. Keller and Mark Monaco 

Background 1 

The Northwestern Hawaiian Islands Biogeographic Assessment 2 

The Region's Unique Natural Environment 6 

Major Taxa of Marine Resources 8 

History of Use and Management 10 

References 12 

Chapter 2: Oceanographic and Physical Setting 17 

Arthur Desch, Timothy Wynne, Russell Brainard, Alan Friedlander and John Christensen 

Introduction 17 

Regional Summary 18 

Ocean Remote Sensing Analysis: Data and Methods 22 

Ocean Remote Sensing Analysis: Results 26 

Existing Data Gaps 39 

Conclusions 39 

Appendix I: Sea Surface Temperature Time Series Plots 41 

Appendix II: Chlorophyll Time Series Plots 51 

References 61 

Chapter 3: Geology and Benthic Habitats 65 

Jonathan Weiss, Joyce Miller, Emily Hirsch, John Rooney Lisa Wedding and Alan Friedlander 

Introduction and Origin 65 

Benthic Habitat Mapping 69 

Existing Data Gaps 101 

References 102 

Chapter 4: Benthic Communities 105 

James Maragos, Jean Kenyon, Greta Aeby Peter Vroom, Bernardo Vargas-Angel, Russell Brainard, 
Lisa Wedding, Alan Friedlander, Jacob Asher, Brian Zgliczynski and Daria Siciliano 

Introduction 105 

Corals 105 

Algae 139 

Invertebrates 145 

Existing Data Gaps 148 

Appendix 150 

References 152 

Chapter 5: Fishes 155 

Alan Friedlander, Edward De Martini, Lisa Wedding and Randy Clark 

Biogeography of Fishes 155 

Latitudinal Affinities Among Fishes 161 

General Fish Assemblage Structure 165 

Existing Data Gaps 187 

References 188 



Chapter 6: Marine Protected Species 191 

Charles Littnan, Marie (Chapla) Hill, Stacy (Kubis) Hargrove, Kaylene E. Keller and Angela D. Anders 

Introduction 191 

Cetaceans 192 

Pinnipeds 211 

Marine Turtles 221 

Existing Data Gaps 228 

References 229 

Chapter 7: Seabirds 235 

Kaylene E. Keller, Angela D. Anders, Scott A. Shaffer, Michelle A. Kappes, Beth Flint and Alan Friedlander 

Introduction 235 

Procellariiformes (Albatrosses, Petrels and Shearwaters) 240 

Pelecaniformes (Boobies, Frigatebirds and Tropicbirds) 249 

Charadriiformes (Terns and Noddies) 256 

Population Status and Trends 263 

Existing Data Gaps 271 

References 272 

Chapter 8: Nonindigenous and Invasive Species 275 

Kevin See, Scott Godwin and Charles Menza 

Introduction 275 

Marine Algae 278 

Invertebrates 280 

Fishes 285 

Management 287 

Existing Data Gaps 287 

References 288 

Chapter 9: Connectivity and Integrated Ecosystem Studies 291 

Alan Friedlander, Donald Kobayashi, Brian Bowen, Carl Meyers, Yannis Papastamatiou, Edward DeMartini, 
Frank Parrish, Eric Treml, Carolyn Currin, Anna Hitting, Jonathan Weiss, Chris Kelley Robert O'Conner, 
Michael Parke, Randy Clark, Rob Toonen and Lisa Wedding 

Introduction 291 

Large-Scale Population Connectivity Models from Ocean Currents 291 

Essential Fish Habitat 316 

Trophic Relationships: Stable Isotope Composition of Primary Producers 

and Consumer Organisms 320 

Food Web Models 324 

Existing Data Gaps 326 

References 328 

Chapter 10: Management Concerns and Responsibilities 331 

Kaylene E. Keller, Angela D. Anders, Ann Mooney Randall Kosaki, Malia Chow and Mark Monaco 

Introduction 331 

Background 331 

World Heritage Nomination 333 

Management of Protected Species 334 

Management of Threats to the Ecosystem 336 

Management of Human Impacts 349 

Monument Permit Applications and Permit Issuance 351 
Future Directions and Implications for a Biogeographic Assessment to 

Support Hawaiian Archipelago Spatial Management 359 

References 361 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

EXECUTIVE SUMMARY 



The mission of NOAA's Office of National Marine Sanctuaries (ONMS) is to serve as the trustee for a system of 
marine protected areas, to conserve, protect and enhance biodiversity. To assist in accomplishing this mission, 
the ONMS has developed a partnership with NOAA's Center for Coastal Monitoring and Assessment's Bioge- 
ography Branch (CCMA-BB) to conduct biogeographic assessments of marine resources within and adjacent 
to the marine waters of NOAA's National Marine Sanctuaries (Kendall and Monaco, 2003). 

Biogeography is the study of spatial and temporal distributions of organisms, their associated habitats, and 
the historical and biological factors that influence species' distributions. Biogeography provides a framework to 
integrate species distributions and life history data with information on the habitats of a region to characterize 
and assess living marine resources within a sanctuary. The biogeographic data are integrated in a Geographi- 
cal Information System (GIS) to enable visualization of species' spatial and temporal patterns, and to predict 
changes in abundance that may result from a variety of natural and anthropogenic perturbations or manage- 
ment strategies (Monaco et al., 2005; Battista and Monaco, 2004). 

Defining biogeographic patterns of living marine resources found throughout the Northwestern Hawaiian Is- 
lands (NWHI) was identified as a priority activity at a May 2003 workshop designed to outline scientific and 
management information needs for the NWHI (Alexander et al., 2004). NOAA's Biogeography Branch and the 
Papahanaumokuakea Marine National Monument (PMNM) under the direction of the ONMS designed and 
implemented this biogeographic assessment to directly support the research and management needs of the 
PMNM by providing a suite of spatially-articulated products in map and tabular formats. The major findings of 
the biogeographic assessment are organized by chapter and listed below. 

















The NWHI are home to a wide variety of ecosystems and living marine resources. Photos: J. Maragos. 

Oceanography 

• Sea surface temperature (SST) analyses suggest three latitudinal subunits: 

North - Kure, Midway, Pearl and Hermes - temperature range 20-27 °C, among the largest 

variation in any coral reef ecosystem; 
Middle - Lisianski to Gardner; and 
Southern - French Frigate Shoals to Nihoa, average temperature range 23-27 °C. 

• SST analysis suggest there may have been bleaching events in 1987 and 1991 based on temperature 
anomalies >1 °C during warm period (August-September). 

• Sea surface fronts or eddies are active in the northern region from December to April with a peak in 
March. Few fronts are found in the southern portion and during mid-summer, fronts retreat to the north. 

• Low productivity ocean water is increasing worldwide due to climate change and this water has reached 
the Hawaiian archipelago with major implications for ecosystem productivity. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Geology and Benthic Habitats 

• Only 50% of shallow water habitats (0-30 m) have been mapped with more than one-half of this habitat 
consisting of hardbottom. 

• Nearly complete high resolution bathymetric coverage from 3 to 3,000 m exists for Kure, Midway, Pearl 
and Hermes, Brooks Bank and French Frigate Shoals. 

• Upchain islands (Kure, Midway, Pearl and Hermes and Lisianski) have steeper slopes compared to down- 
chain islands. The highest rugosity (complexity) derived from multibeam data was at Pearl and Hermes. 

Benthic Communities 

• There are approximately 80 morphologically distinct coral "species" in the NWHI, and about 35 are likely 
to be endemic (44%). This represents some of the highest endemism of any coral assemblage found 
on earth. More than 25 NWHI species are still undescribed or unidentified, and once type specimens 
are collected and examined morphologically and genetically, then final determinations can be made on 
which corals are new species and possible endemics. Notwithstanding efforts to date, NWHI explorations 
are still inadequate, and it is likely that additional undescribed coral species will be encountered in the 
future. 

• The highest species richness was observed at French Frigate Shoals and reflects optimal conditions for 
coral growth (optimal temperature, low wave exposure, large open atoll) 

• Coral cover, based on tow board data, was 8% overall. Lisianski (19%) and Maro (15%) had the highest 
cover while Midway (2%) had the lowest. Coral cover on hard bottom only was highest at Maro (39%) 
and Lisianski (37%). 

• No significant differences were found in coral cover at permanent locations between 2002 and 2006. 

• Disease prevalence is low overall, <1%. 

• Algal diversity is similar across the chain but brown algae are more abundant at the northern end of the 
chain (Midway and Kure). Algal endemism was calculated at 11% for French Frigate Shoals, and 7% for 
Gardner Pinnacles, however recent molecular evidence coupled with more detailed morphological obser- 
vations are finding that a large percentage of the Hawaiian algal flora are likely incorrectly identified and 
may be species new to science. 

Fishes 

• Endemism based on numbers is 52% and 21% based on species richness. 

• Fish assemblages showed latitudinal affinities with more subtropical and temperate species found to the 
north and more tropical species to the south. The major faunal break occurs around Maro and Laysan. 

• Midway and Kure have a distinct fish assemblage based on biomass as does Nihoa and Mokumana- 
mana. These distinct assemblages at the extremes of the chain are likely the result of temperature and 
habitat. 

• Grey reef sharks are more common downchain while Galapagos sharks are more common to the north. 
The transition occurred around Gardner and Maro. 

• Ranking of fish assemblages (based on endemism, species richness, number of individuals, biomass, 
apex predator biomass and recruitment) show Pearl and Hermes with the highest rank, followed by Mid- 
way, and French Frigate Shoals, respectively. 

Protected Species 

• Fifteen cetacean species have been observed within the Monument boundaries. This group of organisms 
has not been well studied in the NWHI and it may represent an important area for these species. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

• Hawaiian monk seals have shown dramatic declines since the late 1990s but these declines differ among 
locations and for various reasons. French Frigate Shoals currently supports the largest colony but has 
also shown the steepest decline (75% since 1989) with low juvenile survival. Laysan and Lisianski have 
remained fairly constant over the past decade. Pearl and Hermes is the only location to have shown an 
increase in the past 10 years. Low population numbers at Midway and Kure are likely the result of long 
histories of human disturbance. 

• Green turtle populations continue to increase in the NWHI, representing a major conservation success 
story. French Frigate Shoals accounts for over 90% of the nesting population with Laysan, Lisianski, and 
Pearl and Hermes accounting for the remainder. Despite extensive potential beach nesting habitat at 
Midway and Kure, limited nesting has been observed at Midway and none at Kure. 

Seabirds 

• Twenty-two species of seabirds exist in the NWHI, representing one of the most important locations for 
seabirds in the tropics. Despite the importance of this region, long-term data collection has been restrict- 
ed to a few species at a limited number of locations. 

• Foraging areas range from only a few kilometers for some species (e.g. Little Terns) to over 1,000 km for 
species like the Laysan Albatross. Wide ranging species will require cooperative management at national 
and international levels. 

• Changes in SST and primary production will result in changes in seabird populations over time and moni- 
toring of these populations will be important to better understand the impacts of climate change. 

Nonindigenous and Invasive Species 

• The presence of invasive species is currently concentrated in harbors and when man-made objects are 
present. 

• The blueline snapper (Taape) have spread from Oahu in 1958 to Midway in 1992, a distance of 1,180 nmi 
and observations at other islands in the chain show during this time period show a rate of spread of about 
18-70 nmi/yr. On the other hand the Peacock grouper (Roi) has only made it as far as French Frigate 
Shoals or about 5-17 nmi/year. 

• The current presence of invasive species is low but data are limited, particularly at deeper depths and 
more systematic sampling is currently being implemented. 

Connectivity and Integrated Ecosystem Studies 

• Advection-diffusion ocean current models show Kure, Midway, and Pearl and Hermes strongly connect- 
ed. Scale of dispersal is on the order of 50-150 km. 

• High-resolution ocean current data and computer simulation showed that for organisms with short larval 
duration (15 days - e.g., corals and some invertebrates), a narrow transitional region from Kauai to Nihoa 
included settlers from both the Main Hawaiian Islands (MHI) and NWHI. For typical fish larvae (45 day lar- 
val duration) there appeared to be little exchange between the NWHI and the MHI north of French Frigate 
Shoals. For the few organisms with longer larval durations (eg., Bottomfishes, lobster) nearly all regions 
of the MHI have at least some NWHI settlers where as most of the NWHI is self-seeding. 

• Giant trevally (ulua) and jobfish (uku) show no inter-atoll movement over a several year period. They con- 
gregate around channels and reef passes and make seasonal, within-atoll movements to spawn. 

• Genetic connectivity for opihi, the Hawaiian endemic limpet, between the MHI and NWHI is too low (less 
than three migrants per generation) to replenish MHI stocks and one species is not even found in the 
NWHI. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

• Barriers to genetic dispersal occur between Pearl and Hermes and Midway, as well as between Niihau 
and French Frigate Shoals for some species however closely related species with similar ecology and 
reproductive biology have dramatically different patterns of connectivity. 

• Potential bottom fish habitat was calculated among islands based on depth (100-400 m), slope (>20%), 
and hardness using available mapping data. Kure (31%) and Maro (30%) had the greatest percentage 
of potential bottom fish habitat to total area mapped. These results may indicate areas with high popula- 
tions and therefore greater replenishment potential for other locations. These results may also be useful 
in identifying monk seal forage habitat. 

• Stable isotope analysis indicates that benthic algae provide the majority of the trophic support for apex 
predators and the entire system consists of short (three to four trophic levels above primary production) 
food chain. These results are very consistent with the Ecopath model estimates of food web supporting 
fisheries production in the NWHI. 



Management 

• The complex meta-population dynamics observed in the NWHI requires a better understanding of how 
these populations replenish themselves and how they connect to other areas. 

• The results show some areas of strong linkage and others with clear breaks in connectivity. This will re- 
quire zoning and other spatial management tools to maintain ecosystem function. 

• Current and future changes in the environment as a result of climate change will require more compre- 
hensive assessment and monitoring along with the ability to respond in a timely manner to mitigate po- 
tential negative impacts to the ecosystem. 

• The data gaps identified in the report (e.g., shallow-water maps, cetaceans, seabirds, deeper habitats, 
etc.) need to be addressed in order to have a better understanding of the entire ecosystem. 

• A biogeographic assessment that integrates the MHI and NWHI would better help to explain ecosystem 
connectivity and processes. 

For questions or more information, please visit http://ccma.nos.noaa.gov/about/biogeography/ or contact: 

Mark E. Monaco, Ph.D. 

Biogeography Branch Chief 

NOAA/NCCOS/CCMA 

1305 East West Highway, N/SCI1 

Silver Spring, MD 20910 

Email: mark.monaco@noaa.gov 

Alan M. Friedlander, Ph.D. 

USGS Hawaii Cooperative Fishery Research Unit 

Department of Zoology 

University of Hawaii 

Honolulu, HI 96822 

Email: alan.friedlander@hawaii.edu 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



REFERENCES 



Alexander, C.A., S.R. Gittings, R. Kosaki, and M.S. Tartt. 2004. Information needs for conservation science and manage- 
ment of the Northwestern Hawaiian Islands: a product of the I ke amio o na wa'a workshop. National Oceanic and Atmo- 
spheric Administration, Marine Sanctuaries Division. 31 pp. & app. 

Battista, T.A. and M.E. Monaco. 2004. Geographic information systems application in coastal marine fisheries. Pages 
189-208 In: W.L. Fisher and R.J. Rahel, editors, Geographic information systems in fisheries. American Fisheries Society, 
Bethesda, MD. 

Kendall, M.S., and M.E. Monaco. 2003. Biogeography of the National Marine Sanctuaries: A Partnership between the 
National Marine Sanctuary Program and the National Centers for Coastal Ocean Science Biogeography Program. Un- 
published Report. 15 pp. 

Monaco, M., Kendall, M., Higgins, J., Alexander, C, and Tartt, M., (2005) Biogeographic assessments of NOAA National 
Marine Sanctuaries: The integration of ecology and GIS to aid in marine management boundary delineation and assess- 
ment, in Wright, D.J. and Scholz, A.J. (Eds.), "Place Matters: Geospatial Tools for Marine Science, Conservation, and 
Management in the Pacific Northwest," Corvallis, OR: Oregon State University Press, 2005. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Introduction 

Alan Friedlander 12 , Kaylene E. Keller 3 and Mark Monaco 1 



BACKGROUND 

The mission of NOAA's Office of National Marine Sanctuaries (ONMS) is to serve as the trustee for a system of 
marine protected areas, to conserve, protect and enhance biodiversity. To assist in accomplishing this mission, 
the ONMS has developed a partnership with NOAA's Center for Coastal Monitoring and Assessment's Bioge- 
ography Branch (CCMA-BB) to conduct biogeographic assessments of marine resources within and adjacent 
to the marine waters of NOAA's National Marine Sanctuaries (Kendall and Monaco, 2003). 



Biogeography is the study of spatial and temporal distributions of organisms, their associated habitats, and 
the historical and biological factors that influence species' distributions. Biogeography provides a framework 
to integrate species distributions and life history data with information on habitats of a region to characterize 
and assess living marine resources within a marine protected area. The biogeographic data are integrated in 
a Geographical Information System (GIS) to enable visualization of species' spatial and temporal patterns and 
to predict changes in abundance that may result from a variety of natural and anthropogenic perturbations or 
management strategies (Monaco et al., 2005; Battista and Monaco, 2004). The complexity of products from 
biogeographic analysis range from simple species distribution maps of a particular habitat, to more complex 
products that combine single data layers to create maps of biodiversity or habitat complexity (NOAA, 2003b; 
Pittman et al., 2007). The biogeographic assessment approach was developed by the CCMA-BB in consulta- 
tion with the ONMS in 2003 (Kendall and Monaco, 2003; Monaco et al., 2005; Figure 1.1). 



The Biogeographic Assessment Approach 



Biogeographic 
Data Layers 



Example Integrated 
Biogeographic Analyses" 



Products to Aid 
Management 



Imagery 



Patterns of 
Human Use 



Bottom Type 



Bathymetry 



Oceanography 



Species 
Distributions 
(many layers) 



S£ Evaluate internal zone 

'S boundaries relative to 

u> biological resources 

5* - 



Species 
Richness 



Threatened 
Habitats 





j2 Explore options for 
2 reducing ecosystem 
— threats 



Evaluate alternative 
management strategies 



Figure 1.1. Generalized biogeographic assessment process developed by CCMA-BB. Source: Kendall and Monaco, 
2003. 



1. NOAA/NOS/NCCOS/CCMA Biogeography Branch 

2. The Oceanic Institute 

3. NOAA/NOS/ONMS/Papahanaumokuakea Marine National Monument 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Typically a biogeographic assessment is comprised of the three primary activities: 1) compile individual bio- 
geographic data layers, 2) perform integrated biogeographic analyses, and 3) develop products to aid in man- 
agement (Figure 1.1). A key tool used to develop and implement the assessment is the use of GIS technology 
which aids in data compilation, spatial analyses, and visualization of results to support place-based manage- 
ment needs (Battista and Monaco, 2004). The assessment process shown in Figure 1.1 is based on geospatial 
and temporal analyses of existing physical and biological data and has resulted in many spatially-oriented 
products that help managers understand how ecosystems function. Often biogeographic analyses focus on 
determining the strength of coupling between habitats and species and defining discrete areas of biological 
significance (NOAA, 2003; NOAA, 2005; Monaco et al., 2005). 

Defining biogeographic patterns of living marine resources found throughout the Northwestern Hawaiian Is- 
lands (NWHI) was identified as a priority activity at a May 2003 workshop designed to define scientific and 
management information needs for the NWHI. NOAA's Biogeography Branch and the Papahanaumokuakea 
Marine National Monument (PMNM) under the direction of the ONMS designed and implemented this bio- 
geographic assessment to directly support the research and management needs of the Monument, such as, 
minimizing impacts of permitted research activities on NWHI marine resources. Successful implementation of 
this assessment required cooperation and participation with many federal, state, academic and private sec- 
tor partners. Without participation of key partners, such as NOAA's Pacific Islands Fisheries Science Center, 
Coral Reef Ecosystem Division, the University of Hawaii, the University of Miami, the State of Hawaii's Division 
of Aquatic Resources, and the U.S. Fish and Wildlife Service, the biogeographic assessment would have not 
been completed. 



THE NORTHWESTERN HAWAIIAN ISLANDS MARINE BIOGEOGRAPHIC ASSESSMENT 

In an effort to provide further protection of the NWHI, the Monument was created by Presidential proclamation 
on June 15, 2006. The Co-Trustees for the Monument are NOAA, the Department of the Interior and the state 
of Hawaii. This biogeographic assessment was designed to support the Monument's scientific and manage- 
ment needs based on historical, recent and planned research and monitoring studies within the Monument 
(Table 1.1). The assessment has resulted in a suite of spatially-articulated products for use by the Monument 

Table 1.1. The monitorinq programs that are currently collecting data in the NWHI. 



MONITORING PROGRAM 



Fishery monitoring and economics 
program 



OBJECTIVES 



Fisheries catch and effort statistics 



YEAR EST. FUNDING 



1948 



NOAA 



AGENCIES 



PIFSC, DAR 



Marine turtle research program 



Monitor selected sea turtle breeding 
sites 



1973 



NOAA, 
USFWS 



USFWS, PIFSC 



Seabird monitoring 



Monitoring selected nesting seabird 
species 



1978 



USFWS 



USFWS, PIFSC 



Fishery independent lobster 
monitoring 



Monitor lobster using fisheries-inde- 
pendent sampling 



1983 



NOAA 



PISSC 



Marine mammal research program Monitor and assess subpopulations 



Marine debris program 

Reef assessment and monitoring 
program 

Oceanography & water quality 
program 



Rates of marine debris accumulation 

Monitor and assess reef communities 
through integrated ecosystem science 

Physical and chemical oceanographic 
conditions and processes influencing 
reef health. 



1985 
1996 

2000 
2000 



NOAA 
NOAA 

CRCP 
NOAA 



PIFSC, USFWS 

CRED, UH, USFWS, 
DAR, USGS 

CRED, USFWS, NMSP, 
DAR, numerous col- 
laborators 



PIFSC-CRED, UH 



Coral monitoring 

Connectivity and ecosystem health 



Monitoring corals at permanent sites 



Examine connectivity, ecosystem 
health, and genetic structure 



2000 



2005 



HCRI, 
USFWS 

NMSP 



USFWS, PIFSC-CRED 



HIMB 



Abbreviatons: CRCP - NOAA's Coral Reef Conservation Program; CRED - Coral Reef Ecosystem Division; DAR - Hawaii 
Division of Aquatic Resources, Department of Land and Natural Resources; HCRI - Hawaii Coral Reef Initiative; HIMB - Hawaii 
Institute of Marine Biology; NMSP - National Marine Sanctuary Program; NOS - National Ocean Service; PIFSC - Pacific Islands 
Fisheries Science Center; UH - University of Hawaii; USGS - U.S. Geological Survey 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

and its partners to support ecosystem-based management and the long-term, comprehensive protection and 
conservation of the marine resources of the NWHI. Results of this assessment will also support the adaptive 
management process, identify gaps in information and direct research priorities. 

Project Objectives 

The biogeographic assessment of the NWHI contributes greatly toward ecosystem-based management of the 
marine resources of the NWHI. The study is broad in scope and includes characterization of the physical and 
biological environments (e.g., oceanography, habitats) that structure the spatial and temporal distribution of 
living marine resources within and adjacent to the Monument boundaries. The objectives of the study were: 

1. Identify and synthesize relevant biological, physical and socioeconomic data sets for the study area. 
Organize the data in a common spatial framework within a GIS. 

2. Conduct a marine biogeographic analysis of available data to identify important ecological linkages and 
biologically significant regions and time periods, based on species distributions, abundance, associated 
habitats and their ecological function. 

The objectives were addressed using the biogeographic assessment process (Figure 1.1) and resulted in a 
synthesis of multiple data sets that range across the NWHI to enable characterization of the biological and 
physical environment that structures the biogeographic patterns in space and time of living marine resources 
found within the Monument. The data, analyses and supporting information are linked using statistical and GIS 
tools to visualize the location of significant biological areas or "hot spots." There were many alternative ap- 
proaches to analyze and organize the biological, physical and habitat data compiled for this assessment. How- 
ever, only a limited number of analytical options were selected based on reviewer's comments on the project 
work plan and technical review meetings. These key analyses are presented in this document. A critical step in 
assessment process was the extensive effort to have data, analytical approaches and results peer reviewed. 
Initial results from the suite of analyses were presented to experts on NWHI marine ecosystem, as well as to 
the originators of the data sources in an attempt to improve the analyses. The role of expert review and input 
has been considerable, and the contributions made by experts have significantly enhanced the study results. 

The use of the GIS enabled species-specific data, such as distribution and abundance data or community 
metrics (e.g., species richness), to be directly linked to specific areas or habitats they correspond with across 
the study area. The GIS also facilitated integration of multiple data types and sources into a common spatial 
and temporal framework (Gill et al., 2001). 

The chapters that follow focus on the spatial and temporal distribution data and analyses from the assessment. 
The report is organized by introducing the geology, habitats and oceanographic characteristics of the NWHI 
and then followed by the biogeography of living marine resource found within and adjacent to the Monument 
(Miller et al., 2003; Maragos et al., 2004). The Connectivity and Integrated Ecosystem Studies chapter ad- 
dresses combinations of the individual biogeographic data layers to characterize the Monument based on the 
integration of physical and biological data, including ecological and genetic connectivity that define the PMNM 
ecosystem. Finally, the Management Concerns and Responsibilities chapter focuses on the management of 
the Monument including the management structure, protected species occurring within the Monument, the 
management of human activities and the greatest potential threats to the region. Below are brief summaries 
that characterize the PMNM ecosystem and are discussed in greater detail in individual sections. 

Region of Interest 

The remoteness and protective status of the NWHI has minimized reef degradation suffered by many other 
coral reefs around the world (Friedlander et al., 2005, 2008; Grigg et al., 2008). The NWHI consist of small is- 
lands, atolls, submerged banks, and reefs, that stretch for more than 2,000 km northwest of the high windward 
Main Hawaiian Islands (MHI; Figure 1.2). The majority of the islets and shoals remain uninhabited, although 
Midway, Kure, Laysan Island and French Frigate Shoals have all been occupied for extended periods over 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Midway 
Kure , Atoll 
Atoll 



\ 



Hermes Atoll 



a S3 



\ 



v 



Lisianski , 
Island 



Laysan 
Island 



\ 



Raita 

Bank Gardner 

Kfl Pinnacles 



X 



2X10 100 km 



French 

Frigate 



Mokumanamana 



- - Papahanaumokuakea MNM 

Exclusive Economic Zone 

] Land 

Water <20 m 

Water <40 m 

| Water <200 m 

Deep Water 




Figure 1.2. The Northwestern Hawaiian Islands, which extend across the north central Pacific, represent a vast, remote 
coral ecosystem that has been subjected to relatively minimal anthropogenic impacts. Map: K. Buja. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

the last century by various government agencies. The inaccessibility, limited fishing and lack of other human 
activities in the NWHI have resulted in minimal anthropogenic impacts (Friedlander et al., 2005, 2008), there- 
fore providing a unique opportunity to assess how a "natural" coral reef ecosystem functions in the absence of 
major localized human intervention and contrast these with anthropogenic influences experienced in the MHI 
and other comparable ecosystems (Friedlander and DeMartini, 2002; Grigg et al., 2008). 

One of the most striking and unique components of the NWHI ecosystem is the abundance and dominance of 
large apex predators such as sharks and jacks (Figure 1.3; Hobson, 1984; Parrish et al., 1985; Friedlander and 
DeMartini, 2002), which exert a strong top-down control on the ecosystem (DeMartini et al., 2005; DeMartini 
and Friedlander, 2006) and have been depleted in most other locations around the world (Meyer and Worm, 
2003; 2005). The NWHI flora and fauna include a large percentage of species that are endemic to the Hawaiian 
Islands, which are recognized for having some of the highest marine endemism in the world (Kay and Palumbi, 
1987; Jokiel, 1987; Randall, 1998; Randall, 2007). Some of these endemics are dominant components of the 
community, resulting in a unique ecosystem that has extremely high conservation value and has identified Ha- 
waii as an important global biodiversity hot spot (DeMartini and Friedlander, 2004; Maragos et al., 2004). The 
few alien species known from the NWHI are restricted to the anthropogenic impacted habitats of Midway Atoll 
and French Frigate Shoals (Friedlander et al., 2005, 2008; Godwin et al., 2006). Disease levels in corals in the 
NWHI are much lower than those reported from other locations in the Indo-Pacific (Aeby, 2006). 
The NWHI represent important habitat for a number of threatened and endangered species. 




Figure 1.3. Apex predators are a conspicuous and important component of the PMNM ecosystem. Galapagos sharks 
(Carcharhinus galapagensis; left) sharks at Maro Reef and giant trevally (Caranx ignobilis; right), known as ulua in Hawaii, 
from French Frigate Shoals. Photos: J. Maragos. 

The Hawaiian monk seal is one of the most critically endangered marine mammals in the U.S. (approximately 
1,200 individuals) and depends almost entirely on the islands of the NWHI for breeding and the surrounding 
reefs for sustenance (Antonelis et al., 2006). Over 90% of all sub-adult and adult Hawaiian green sea turtles 
found throughout Hawaii inhabit the NWHI (Balazs and Chaloupka, 2006). Additionally, seabird colonies in 
the NWHI constitute one of the largest and most important assemblages of seabirds in the world (USFWS, 
2005). 

On June 15, 2006, President George W. Bush designed the NWHI as a Marine National Monument, the largest 
no-take marine conservation areas on earth, through the signing of Proclamation 8031. In March 2007, First 
Lady Laura Bush renamed the Monument the Papahanaumokuakea Marine National Monument on behalf of 
the President. The Monument encompasses nearly 225,300 km 2 of ocean and includes all the islands, atolls, 
shoals and banks from Nihoa Island to Kure Atoll (Figure 1.4). The unique predator-dominated trophic struc- 
ture, the dominance by large numbers of endemic species, and the occurrence of a number of threatened and 
endangered species makes the NWHI an ecosystem of global significance. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Papahanaumokuakea 
Marine National Monument 



V 'Eii FfearlandX 
\ X_ J Hermes Atoll X 

^-— . P K> 


— - — . SJmai* ^V 






^V fc/V*' 


Legend \^ 


~j Marine National Monumen 


t Boundary 


H 100 Fathom Contours 




Special Preservation Area 

100fm: Kure Atoll, Pearl and Hermes Aloll 

5Wrn: Laysan Island 

25fm: Lrsianski Island. Maro Reef. Gardner Pinnacles, Meeker Island 

3nm: Nihoa 

French Frigate 5noaJS BOuntfaTy COWdlnalflS-; 

(166* 45' W. 24' 10' NH165* 35' W, 24* N> 

( 1 66' AS W, 24' T NJ (1 W 55' W.. 23' 41 ' N) 

(IBS- 55' W, 24* 2' N) (163* 35 W, 29* 30' N) 

Ecological Reserves 

Existing Commercial Fishing Area Phased Out by June 2011 

Managed as Ecofogtcaf Reserve fofkrmng priase-ouJ 

I Special Management Area 
Boundary txttneh 12nm from ttnd 

1"^ ^ Emergent Land Features 

SOgn» Cffl-i prcvi^cl by NOAA. Si3fq <?T Hawaii arid ESHI 



Figure 1.4. Papahanaumokuakea Marine National Monument boundaries. Map: PMNM. 



THE REGION'S UNIQUE NATURAL ENVIRONMENT 

The NWHI are influenced by a dynamic environment that includes large annual water temperature fluctuations, 
seasonally high wave energy, and strong inter-annual and inter-decadal variations in ocean productivity (Grigg, 
1983; Polovina et al., 2001). As a result of these influences and the general absence of human interference, 
natural stressors play an important role in the structure of the NWHI ecosystem (Friedlander et al., 2005). 
Large swell events generated every winter commonly produce waves up to 10-12 m in height, which limits the 
growth and abundance of coral communities, and leads to species and growth forms that are adapted to these 
dynamic high wave energy environments (Grigg et al., 2008). 

Compared with most reef ecosystems around the globe, the large annual fluctuation of sea surface tempera- 
tures (SSTs; >10°C) found at the northernmost atolls of the NWHI is extremely high. Cooler water temperatures 
to the north restrict the growth and distribution of a number of coral species (Grigg, 1983), and the biogeo- 
graphic distribution of many fish species in the NWHI is influenced by differences in water temperatures along 
the archipelago (DeMartini and Friedlander, 2004; Mundy, 2005). 



Large-Scale Biogeographic Regions 

The NWHI is set in a dynamic oceanographic and meteorological regime in the northern/central subtropical 
region of the Pacific Ocean (Figure 1.5). The NWHI archipelago extends such a great distance that the two 
opposite ends of the chain often experience somewhat different oceanographic and meteorological conditions 
(Table 1.2). The NWHI are not usually impacted by tropical storms but do experience large boreal winter wave 
events that assist in shaping the ecosystem. The boundary between the nutrient-poor, or oligotrophic, surface 
waters of North Pacific Subtropical Gyre and the nutrient-rich, or eutrophic, surface waters of the North Pacific 
Subpolar Gyre is frequently in the NWHI region (Leonard et al., 2001). This front shifts 15° (between 30° and 
45°N) seasonally (Polovina et al., 2001), reaching far enough south in the winter to encompass the northern 
most three atolls (Kure, Midway, and Pearl and Hermes). The southern extension of this front into the NWHI re- 
gion, migrating on interannual and decadal time scales, brings colder and nutrient rich waters that are likely im- 
portant to the productivity and ecology of these coral reef ecosystems. The location of this front also influences 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



marine debris concentrations (Kubota, 
1994), which has been shown to most 
severely impact the northern atolls 
(Boland et al., 2004; Donohue et al., 
2001). 



Geology and Evolutionary History 
The Hawaiian Archipelago originated 
over a relatively stationary melting 
anomaly or hot spot in earth's mantle 
located below the floor of the Pacific 
Plate, which is drifting over the hot spot 
to the northwest at a rate of about 8 cm/ 
yr (Figure 1.6; Griggetal.,2008; Rooney 
et al., 2008). Because the plate is slowly 
cooling as it moves away from the hot 
spot, the overriding islands and other 
volcanic features are slowly subsiding. 
Beginning at Nihoa and Mokumana- 
mana Island (approximately seven and 
10 million years old, respectively) and 
extending to Midway and Kure Atolls 
(approximately 28 million years old), the 
NWHI represent the older portion of the 
emergent Hawaiian Archipelago (Grigg, 
1988; Juvik and Juvik, 1998; Rooney et 
al., 2008). 

The Hawaiian Archipelago is 3,900 km 
from the west coast of the U.S. mainland, 
3,800 km from Japan, and over 1,000 km 
from the nearest island archipelago (the 
Line Islands). This geographic isolation 
of the entire Hawaiian Island chain has 
resulted in a large number of species 
that are found nowhere else on earth, 
and the proportion of these endemic 
species (>25% for most taxa) to other 
native species is some of the highest re- 
corded for any tropical marine ecosys- 
tem to date. Some of these endemics 
are dominant components of the coral 
reef community, resulting in a unique 
ecosystem that has extremely high con- 
servation value (DeMartini and Fried- 
lander, 2004; Maragos et al., 2004). 




Figure 1.5. Topographic map showing location in Pacific Ocean of the 
Northwestern Hawaiian Islands (NWHI) and the major ocean currents in the 
region (North Equatorial Current (NEC, South Equatorial Current (SEC), 
North Equatorial Counter Current (NECC), South Equatorial Counter Cur- 
rent (SECC), Equatorial Under Current (EUC). Source: CRED. 

Table 1.2. Coordinates and Hawaiian names of the 10 islands presented 
from north to south. 



ISLAND HAWAIIAN NAME LATITUDE LONGITUDE 


Kure Atoll 


Kanemilohai, „ QO oc ,. . 
Mokupapapa 28 26 N 


178° 19'W 


Midway Atoll 


Pihemanu 28° 14'N 


177° 22'W 


Pearl and Hermes Atoll 


Holoikauaua 27° 51'N 


175° 51'W 


Lisianski Island and 
Neva Shoals 


Papaapoho 26° 3'N 


173° 58'W 


Laysan Island 


Kauo 25° 47'N 


171° 44'W 


Maro Reef 


Nalukakala, Koanakoa 25° 27'N 


170° 37'W 


Gardner Pinnacles 


Puhahonu 24° 52'N 


168° l'W 


French Frigate Shoals 


Mokupapapa, Kanemilohai 


23° 46'N 


166° 12'W 


Mokumanamana Island 


Mokumanamana 


23° 27'N 


164° 31'W 


Nihoa Island 


Nihoa 


23° 5'N 


161° 51'W 





Figure 1.6. Lava flowing into the ocean off the southeast coast of the island 
of Hawaii. Photo: Hawaii Volcano Naitonal Park. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

MAJOR TAXA OF MARINE RESOURCES 

Benthic Communities 

The composition and distribution of benthic communities in the NWHI reflect the interaction of numerous influ- 
ences, including isolation, latitude, exposure, and the successional age of the island they inhabit. Mosaics of 
habitats range from coral-dominated areas to vast expanses of unconsolidated sediments such as sand and 
mud inhabited by a few epi-benthic fauna. 

Despite their high latitude location, slightly more species of coral have been reported from the NWHI (52 spe- 
cies) compared with the MHI (48 species; Maragos et al., 2004), although a number of potentially new species 
have recently been discovered (Friedlander et al., 2008). Kure is the world's most northern atoll (28° 26'N) and 
is referred to as the Darwin Point, where coral growth and subsidence /erosion balance one another (Grigg, 
1982). Beyond this point lies a chain of seamounts that get progressively deeper and older as they move away 
from a stationary melting point or hot spot in the Pacific plate located southeast of the big island of Hawaii 
(Grigg, 1988). 

The coral assemblage in the NWHI con- 
tains a large number of endemics (ap- 
proximately 30%), including at least sev- 
en species of table corals (acroporids), 
which are the dominant reef-building 
coral in the Indo-Pacific, but are absent 
from the MHI (Figure 1.7, Maragos et al. 
2004). Coral disease is currently low in 
the NWHI; but increases in the frequen- 
cy and intensity of bleaching events due 
to global warming could stress corals 
and make them more susceptible to dis- 
ease. 

Unlike the MHI where alien and invasive 

algae have overgrown many coral reefs, 

the shallow reefs in the NWHI appear 

to be free of invasive algae, and high 

natural herbivory (grazing) results in a 

pristine algal assemblage. Algal diversity appears similar across the NWHI chain even though brown algae 

tend to be more abundant at Midway and Kure Atolls than at most other islands (Vroom and Page, 2006). The 

cooler SSTs found at Kure and Midway Atolls during winter months may favor a higher abundance of brown 

algal species. Lower abundance of green algae at Midway may result from higher herbivorous fish densities 

at this atoll system, suggesting possible top-down control of the benthic habitat (DeMartini and Friedlander, 

2004, 2006). 

The NWHI are documented to contain the highest percent cover of algal species when compared to other 
geographic locations throughout the U.S. tropical Pacific, and the lowest percent cover of living coral (Vroom 
et al., 2006). This is likely due to the subtropical location of the NWHI and cool SSTs that bathe biological 
communities during winter months. Despite relatively high algal populations, the NWHI remain a healthy and 
thriving marine ecosystem. 




Figure 1.7. Table coral (Acropora cytherea) at French Frigate Shoals. Pho- 
to: J. Watt. 



Fishes 

A total of 457 reef and shore fishes have been reported from the MHI while 258 are documented from Midway 
Atoll in the NWHI (Randall et al., 1993). Despite these differences, the total number of species observed on 
quantitative transects in the NWHI (210) and was similar to the 215 species reported in a recent comprehen- 
sive quantitative study around the MHI (Friedlander et al., 2005). The lowest overall fish species richness in 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

the NWHI occurs at the small basalt islands (Mokumanamana, Gardner, Nihoa). It is highest at French Frig- 
ate Shoals and Pearl and Hermes Atoll. Greater species richness at French Frigate Shoals may be related to 
higher coral richness and greater habitat diversity (Maragos et al., 2004) while large area, habitat diversity and 
the presence of subtropical and temperate species that occur at greater depths influence greater richness at 
Pearl and Hermes. 

Reef fish trophic structure in the NWHI is strongly dominated by carnivores (Hobson, 1984; DeMartini and 
Friedlander, 2006). Fish biomass in the NWHI is nearly three times greater than in the MHI with most of the 
difference resulting from large apex predators (primarily sharks and jacks; Friedlander and DeMartini, 2002). 
Today, these top carnivores, analogous to lions and wolves on land, are seldom encountered by divers in the 
inhabited Hawaiian Islands. A number of species such as the endemic spectacled parrotfish (Chlorurus per- 
spicillatus), the endemic Hawaiian hogfish (Bodianus bilunulatus), and bigeye emperor (Monotaxis grandocu- 
lis) are quite abundant and obtain large size in the NWHI. These species are heavily exploited for commercial, 
subsistence and recreational use in the MHI, and their reduced number and sizes in the MHI is likely the result 
of years of chronic overfishing (Friedlander and DeMartini, 2002). 



Seabirds 

Seabird colonies in the NWHI constitute 
one of the largest and most important 
assemblages of seabirds in the world, 
with approximately 14 million birds (5.5 
million breeding annually) representing 
22 species (Friedlander et al., 2005; 
USFWS, 2005). More than 95% of the 
world's Laysan albatross (Phoebas- 
tria immutabilis; Figure 1.8) and Black- 
footed albatross (P. nigripes) nest in the 
NWHI (USFWS, 2005). For several oth- 
er species such as Bonin Petrel (Ptero- 
droma hypoleuca), Christmas Shear- 
water (Puffinus nativitatis), Tristram's 
Storm-petrel (Oceanoframa tristrami) 
and Grey-backed Tern (Sterna lunata), 
the NWHI supports colonies of global 
significance. The last complete inven- 
tory of NWHI breeding populations was 
done between 1979 and 1984 (Fefer et 

al., 1984). Population trends since then have been derived from more intensive monitoring at three islands 
and are stable or increasing for most species at these locations but there is concern for a few, especially the 
albatross due to ingestion of plastics, loss of forage base and other large-scale issues (USFWS, 2005). 




Figure 1.8. Laysan Albatrosses at Midway Atoll NWR. Photo: A. Friedland- 
er. 



Protected Species 
Hawaiian Monk Seal 

The Hawaiian monk seal (Monachus schauinslandi) is listed as Endangered under the Endangered Species 
Act (ESA) and Depleted under the Marine Mammal Protection Act. It is the only endangered pinniped occurring 
entirely within U.S. waters. Monk seals occur throughout the Hawaiian Archipelago, and although most are 
found in the NWHI, a small but increasing number pup in the MHI. They commonly occur on isolated beaches 
for resting, molting, birthing and nursing offspring, spending nearly two-thirds of their time in marine habitats 
(Antonelis et al., 2006). A recent monk seal recovery report estimates that there are only 1,200 Hawaiian monk 
seals still alive (Figure 1.9) and if the declining population trend continues there will be fewer than 1,000 within 
the next three or four years, a decrease of more than 60% since the 1950s. When compared historically, the 
monk seal beach count abundance index reached record lows in 2005. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Hawaiian Green Sea Turtle 
The green sea turtle (Chelonia mydas) 
is the most abundant large marine her- 
bivore globally (Bjorndal, 1997) and has 
a circumtropical distribution with distinct 
regional population structures (Figure 
1.10; Bowen et al., 1992). Worldwide, 
the green turtle has been subject to a 
long history of human exploitation with 
some stocks now extinct and others in 
decline. Green sea turtles in U.S. waters 
have been protected under the federal 
ESA since 1978. The Hawaiian green 
turtle stock, or honu, comprises a single 
closed genetic stock that is endemic to 
the Hawaiian Archipelago (Bowen et 
al., 1992) with numerous distinct forag- 
ing grounds within the 2,200 km span 
of the Hawaiian Archipelago From the 
mid-1800s until about 1974, the Hawai- 
ian stock was subject to human exploi- 
tation such as turtle harvesting at forag- 
ing grounds, harvesting of nesters and 
eggs, and nesting habitat destruction. 




Figure 1.9. Hawaiian monk seal and endemic Hawaiian green sea turtle at 
NWHI. Photo: National Marine Fisheries Service. 




Figure 1.10. Endemic Hawaiian green sea turtle at Midway Atoll. Photo: J. 
Watt. 



The primary rookery for the Hawaiian 
green sea turtle is located on French 
Frigate Shoals which accounts for more 

than 90% of all nesting within the Hawaiian Archipelago. The main rookery island at French Frigate Shoals 
is East Island where at least 50% of all French Frigate Shoals nesting occurs. Nesting females exhibit strong 
island fidelity, and the Hawaiian green sea turtle stock has been continuously monitored for several decades. 
Annual surveys of the number of female green turtles coming ashore to nest each night have been conducted 
at East Island since 1973 (Balazs, 1980). 

The long-term trends based on a population model for the East Island nester abundance illustrates a dramatic 
increase in abundance over the past 30-years, and substantial fluctuations in the number of annual nesters 
has been observed (Balazs and Chaloupka, 2006). Such fluctuations are characteristic of green turtle nesting 
populations and reflect a variable proportion of females in the population that breed each year in response to 
ocean-climate variability. The Hawaiian green sea turtle stock is showing signs of recovering after more than 
25 years of protecting their nesting and foraging habitats in the Hawaiian Archipelago (Chaloupka and Balazs, 
2007). 



HISTORY OF USE AND MANAGEMENT 

The designation of the Monument is the most recent increase in protections and management of the NWHI. 
The NWHI has a long history of human use and increasing efforts for conservation management (Figure 1.11). 
Native Hawaiians traversed and seasonally used the NWHI for hundreds of years and continue today to main- 
tain their strong cultural ties to the land and sea of the NWHI. Post-Western-contact, the NWHI continued to be 
explored and the harvest of natural resources including guano mining, egg harvesting, fishing and other natu- 
ral resource extraction occurred. The U.S. Military maintained active military bases during much of the 1900s. 
In 1909 the first natural resource protection was put in place with the designation of the Hawaiian Islands Bird 
Reservation. Since that time, additional reserves and refuges have been established to protect this unique 
ecosystem. The most recent protections were implemented with the designation of the Papahanaumokuakea 
Marine National Monument June 15, 2006. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Archeological remains show evidence of places of 
worship and home sites on Mokumanamana and 
Nihoa for at least a 500-700 hundred year period 
(Emory, Cleghorn 1988, Irwin 1992). 



10( 



Commercial harvest of whales, seals, turtles, sharks, 
see cucumbers, and pearl oysters by European 
explorers 



18 )s 



1£ 2 



Laysan and Lisianski claimed by the Kingdom of 
Hawaii 



l 57 



U.S. takes formal possession of Midway Atoll. 



1 39 



It 6 



is 3 Transpacific cable station in operation at Midway. 



President Theodore Roosevelt signs Executive Order 
to include NWHI in National Wildlife Refuge System. 



l )9 



Tanager Expedition - Collected biological samples 
and documented changes at Laysan Islands from 
guano mining and resource harvest in the 19th and 
early 20th century. 



19 



French Frigate Shoals becomes a U.S. Naval air 
facility 



is 



Battle of Midway - 100-200 mi north of Midway. This 
decisive victory was the turning point for the war in 
the Pacific. 



Mid 1970 
early 



19£ s 



First NWHI Rapid Assessment and Monitoring 
Program (RAMP) cruise. Multi-disciplinary, multi- 
agency investigation of biological, physical, 
chemical, and geological characteristics of the NWHI. 



1 



J 



President George W. Bush issues Presidential 
Proclamation establishing Northwestern Hawaiian 
Islands Marine National Monument 



1 



J 



IYB 



Seabirds feathers and eggs taken by Japanese and 
under permit from the Kingdom of Hawaii 



l >3 



IE 9 



V 

0s 



r 



2C )0 



2C )6 



Nihoa claimed under the Kamehameha Monarchy 



Kure Atoll annexed by King Kalakaua. 



3s Pan American Clippers fly from Honolulu to Midway 



U.S. Navy begins construction of Pacific Naval Air 

Station 



Cooperative Quadripartite Program. NWHI fishery 

investigations involving National Marine Fisheries 

Service, U.S. Fish and Wildlife Service, Hawaii 

Division of Aquatic Resources, and the University of 

Hawaii. 



President Clinton signs Executive Order creating the 
NWHI Coral Reef Ecosystem Reserve. 



Figure 1.11. Time line of the history of the NWHI use and management. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Oceanographic and Physical Setting 

Arthur Desch 1 , Timothy Wynne 1 , Russell Brainard 2 , Alan Friedlander 34 and John Christensen 1 



INTRODUCTION 

The Northwestern Hawaiian Islands 
(NWHI) are set in a dynamic oceano- 
graphic and meteorological regime in 
the northern/central subtropical region 
of the Pacific Ocean (Figure 2.1). The 
boundary between the nutrient-poor sur- 
face waters of North Pacific Subtropical 
Gyre and the nutrient-rich surface wa- 
ters of the North Pacific Subpolar Gyre 
frequently influence the NWHI region 
(Kazmin and Rienecker, 1996; Leonard 
et al., 2001; Polovina et al., 2001). This 
front shifts seasonally (Polovina et al., 
2001) and migrates on interannual and 
decadal time scales, bringing colder and 
nutrient rich waters that are likely impor- 
tant to the productivity and ecology of 
the region (Polovina and Haight, 1999; 
Nakamura and Kazmin, 2003; Polovina, 
2005). Longer-term changes, partiCUlar- 






Peprl & Hermes 
Atoll . 

L.sianski __ |_aysan. 
\ Island | S | 3fld 





Papahanaumokuakea 
Marine National Monument 



D'ahu 

ilt Motoka "' 



400 Kilometers 



A 



Figure 2.1. Hawaiian Archipelago Including the NWHI (Nihoa Island to Kure 
Atoll) and Main Hawaiian Islands! (Hawaii to Kauai). Inset shows the Ha- 
waiian Archipelago in the Pacific Ocean. Source: PMNM, 2008. 



ly those related to climate, are of concern since the reef ecosystems of the NWHI may not have encountered 
such conditions for hundreds, thousands or even millions of years (Rooney et al., 2008). 

The health, functioning and biogeography of ecosystems of the NWHI are primarily controlled by the oceano- 
graphic processes and conditions, both physical and chemical, to which they are exposed. The Monument's 
diverse biological ecosystems, including fishes, corals and other invertebrates, algae, turtles, seabirds and 
marine mammals, is significantly influenced by ocean currents, waves, nutrients, temperature, and other mea- 
sures of water quality and oceanographic conditions. The most important factors controlling the distribution 
and abundance of coral reefs in the NWHI are depth and shelter from large open ocean winter swell (Grigg. 
1983). 

This chapter provides a comprehensive analysis of ocean currents, waves, temperature, winds and productivity 
using satellite remote sensing data to offer a quantitative assessment of regional ocean climate. The objective 
is to capture spatial and temporal patterns in each of these parameters and to set context for the biogeographic 
assessment that follows. The Monument sits in a region of the Pacific Ocean that is dominated by large-scale 
circulation patterns that fluctuate over periods of years and decades. With that perspective, these data sets 
have been collected and analyzed over a spatial scale that includes much of the North Pacific Ocean. Large 
scale events and processes are the focus here. To illustrate this concept, Figure 2.2 provides a view of global 
sea surface temperature (SST), chlorophyll (ocean color), wind and sea surface height or SSH (proxy for cur- 
rents). Each of these examples represents the average condition, worldwide, for the month of March. This view 
is provided as a reminder that while the NWHI cover a vast region, variability in oceanographic conditions as 
resolved by remote sensing platforms is most evident at basin-wide scales. That is, analysis of remote sens- 
ing data within the NWHI is not as variable as the Pacific basin as a whole. A short summary of regional cli- 



1 . NOAA/NOS/NCCOS/CCMA Coastal Oceanographic Assessment Status and Trends Branch 

2. NOAA/NMFS/Pacific Islands Fisheries Science Center, Coral Reef Ecosystem Division 

3. NOAA/NOS/NCCOS/CCMA Biogeography Branch 

4. The Oceanic Institute 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



mate and oceanography is presented 
in advance of the quantitative analysis 
to provide points of reference for the 
results presented. 



REGIONAL SUMMARY 

Climate 

The climate of the entire Hawaiian Ar- 
chipelago features mild temperatures 
year-round, moderate humidity, per- 
sistent northeasterly trade winds and 
infrequent severe storms. Hawaii's 
climate is notable for its low day-to- 
day and month-to-month variability 
(Giambelluca and Schroeder, 1998). 
The climate is influenced by the ma- 
rine tropical or marine Pacific air 
masses depending upon the season. 
During the summer, the Pacific High 
Pressure System dominates, with the 
ridge line extending across the Pa- 
cific north of Kure and Midway. This 
places the region under the influence 
of easterly winds, with marine tropi- 
cal and trade winds prevailing. During 
the winter, especially from November 
through January, the Aleutian Low 
moves southward over the North Pa- 
cific, displacing the Pacific High (Grigg 
et al., 2008). The Kure-Midway region 
is then affected by either marine Pa- 
cific or marine tropical air, depending 
upon the intensity of the Aleutian Low 
or the Pacific High Pressure System 
(Amerson et al., 1974). The surround- 
ing ocean has a dominant effect on 
the weather of the entire archipelago. 
Air temperature at the northern end of 
the archipelago varies between 11 and 
33°C. Air temperature measurements 
made at six sites on Nihoa Island (23° 
N latitude) from March 2006 to March 
2007 ranged between 16 and 34°C. 
Annual rainfall over the last 26 years 
has been 73.28 cm on average, rang- 
ing between 40.61 and 104.24 cm/ 
year (PMNM, 2008). 




133" 180 








-'ieo° -135° 




Figure 2.2. (A) Global climatological SST (°C), (B) chlorophyll mg/m 3 , (C) 
wind m/sec (D) and sea surface height anomaly cm estimates for the month 
of March. The NWHI study area is shown as a black box. 



El Nino - Southern Oscillation (ENSO) and Pacific Decadal Oscillation (PDO) 

ENSO is an interannual global climate phenomenon that results from the large-scale coupling of atmospheric 
and oceanic processes, which creates significant temperature fluctuations in the tropical surface waters of the 
Pacific and other oceans. The two distinct ENSO signatures in the Pacific Ocean are known as El Nino and La 
Nina. During El Nino events, the Aleutian Low pressure system tends to be more intense and extend further to 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Figure 2.3. Diagram SST anomalies in November 2007 showing La Nina 
conditions. Blue tones indicate cooler than average surface temperatures, 
while orange tones indicate warmer than average. The Hawaiian Islands, 
including the NWHI are in the black box. 



the south (closer to the NWHI), thereby 
producing stronger winds, larger waves 
and cooler water temperatures in the 
NWHI (Bromirski et al., 2005). During 
La Nina, SSTs in the eastern tropical 
Pacific are below average, and temper- 
atures in the western tropical Pacific are 
above average (Figure 2.3). Leonard 
et al. (2001) and Rooney et al. (2008) 
have suggested that positive ENSO 
signatures (warming) correspond with 
southern extensions of the North Pa- 
cific subtropical front. A strong band of 
cool water (blue in false color range) 
appears along the Equator, particularly 
strong near South America. Warm con- 
ditions (orange in false color range) ap- 
pear north and south of this strong blue 
band (Figure 2.3). The NWHI be seen 
straddling both warm and cold portions 
of the basin-wide temperature anomaly, 

highlighting complex regional thermal structure. Because biological communities are significantly influenced 
by spatially and temporally-varying ocean currents, temperature and nutrients (Polovina et al., 1995; Seki et 
al., 2002; Polovina et al., 2004), regional biogeography is equally complex and dynamic. 

The PDO is a long-lived El Nino-like pattern of Pacific climate variability. While the two climate oscillations have 
similar spatial footprints, they have very different behavior in time. Two main characteristics distinguish PDO 
from ENSO are: 1) 20th century PDO events persisted for20-to-30 years, while typical ENSO events persisted 
for 6 to 18 months; and 2) the climatic effects of PDO are most visible in the North Pacific, while secondary 
signatures exist in the tropics. The opposite is generally true for ENSO. Several independent studies provide 
evidence that two full PDO cycles have occurred over the past century, where cool regimes prevailed from 
1890-1924 and again from 1947-1976, and warm regimes dominated from 1925-1946 and from 1977 through 
the mid-1990s. Additional research suggests that 20th century PDO fluctuations were most energetic in two 
general periodicities, one from 15-to-25 years, and the other from 50-to-70 years (http://jisao.washington.edu/ 
pdo/). 

Ocean Temperature 

SST is an important physical factor influencing coral reefs and other marine ecosystems of the Monument. 
Maximum monthly climatological mean SST measured over the last 20 years at Kure is 27 °C in August and 
September (NOAA Pathfinder SST time series; Hoeke et al., 2006), with monthly minimums in February at 19 
°C. The large seasonal temperature fluctuations at the northern end of the archipelago result in the coldest - 
and sometimes the warmest - SSTs in the entire Hawaiian chain (Brainard et al., 2004). At the southern end 
of the Monument, the annual variation in SST is much less, with French Frigate Shoals only varying between 
23.3 and 27.5° C. 



Winter temperatures tend to be 3-7°C cooler at the northerly atolls than at the southerly islands and banks as 
the subtropical front migrates southward. These cooler winter temperatures are thought to reduce coral growth 
rates (Grigg 1983, Grigg et al. 2008). In addition to the strong annual cycle, SST observations show significant 
interannual and decadal variability (Figure 2.4). The highest summer maximum SSTs at the northern atolls oc- 
curred during the summers of 1987, 1991 and 2002, possibly suggesting a teleconnection with ENSO events. 
Winter minimum temperatures at the northern atolls appear to oscillate over a longer time period, as indicated 
by a significant warming of winter SSTs beginning in 1999 and lasting for several years (Brainard et al., 2004). 
During the period between July and September 2002, ocean temperatures along the Hawaiian Archipelago 
were warmer than average. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



■Nihoa 

Gardner Pinnacles 
■Lisianski 
■Kure 



Pearl and Hermes 



French Frigate Shoals 
Laysan 
■Special Management Area (Midway) 



30 




16 



& & £ 
s^ s^ ^ s^ s^ s^ s^ ^ ^ ^ s^ s^ s^ s^ s^ s^ s^ s^ s^ s^ ^ •$? 



<£> <£> $> _& 



& <£> qj* & & <£ 



& & & & 



& & 



& & & 



Figure 2.4. Pathfinder SST (°C ) time-series for islands, atolls and shoals throughout the Monument (1985-2006). 

While coral bleaching can be caused by a wide range of environmental variables acting alone or in combina- 
tion (Jokiel and Brown, 2004), the predominant cause of increasing incidences of coral bleaching globally is 
believed to be persistent warmer than average water temperatures (Jokiel and Coles, 1990; Kenyon et al., 
2006a, b), and indeed a significant bleaching event was documented during the summer of 2002 (Friedlander 
et al., 2005; Hoeke et al., 2006, Kenyon et al., 2006a, b). 

Ocean Currents 

Ocean currents transport and distribute larvae among and between different atolls, islands and submerged 
banks of the NWHI, and also provide the mechanism by which species are distributed to and from the main 
eight Hawaiian Islands, as well as other regions (Polovina et al., 1995). The relatively low species diversity and 
high endemism of the NWHI are the result of the relative oceanographic isolation of the Hawaiian Archipelago 
(Grigg et al., 2008; Demartini and Friedlander, 2004; Friedlander et al., 2008). 

Ocean currents are measured and monitored in the NWHI in many different ways. Since 1990, ocean cur- 
rent profiles along the Hawaiian Archipelago have been measured using Acoustic Doppler Current Profilers 
(ADCP) aboard the NOAA ships Townsend Cromwell (1990 to 2002) and Oscar Elton Sette (2003 to present) 
during routine transects along the archipelago to support a number of scientific cruises for NOAA's Pacific Is- 
lands Fisheries Science Center (PIFSC). 



Based on 10 years of ADCP data (1990-2000), Firing et al. (2004a) demonstrated that upper ocean currents 
in the NWHI are highly variable in both speed and direction. Averaged over time, the resultant mean flow of 
the surface waters tend to flow predominantly from east to west in response to the prevailing northeast trade 
winds. The lack of coral reef ecosystems and low biodiversity to the east, or upstream, of the Hawaiian Ar- 
chipelago explains the low species richness and high endemism (PMNM, 2008). Surface Velocity Program 
current drifters and autonomous profiling explorer drifters have also been deployed in the NWHI by PIFSC 
annually since 2001. These drifters provide indications of the Lagrangian (or water-following) flow, thereby 
representing potential larval pathways (Firing et al., 2004a). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Ocean Waves 

Common throughout the region and perhaps more significant as a natural process affecting the geology and 
ecology of the Monument, are the extra-tropical storms and significant wave events that regularly move across 
the North Pacific in the boreal winter (Friedlander et al., 2008, Grigg et al., 2008). Among each of the islands, 
atolls and submerged banks of the NWHI, the distributions of species of corals and algae, and their associated 
fish and invertebrate assemblages are often determined not only by the ocean currents, but also by the expo- 
sure to ocean waves. Many species of corals and algae can only survive in sheltered or quiescent habitats. 
Other species; however, can survive or even thrive in the high wave-energy habitats on the northwestern fac- 
ing reefs that are exposed to tremendous waves caused by winter storms across the North Pacific. 



These large wave events, greater than 10 m, influence the growth forms and distribution of coral reef or- 
ganisms (Dollar, 1982; Dollar and Grigg, 2004; Grigg et al., 2008) and affect the reproductive performance 
of winter-breeding seabirds nesting on 
low islets in the Monument. Most large 
wave events, 5 to 10 m, approach the 
NWHI from the west, northwest, north 
and northeast, with the highest energy 
generally occurring from the northwest 
sector. The southern sides of most of 
the islands and atolls of the NWHI are 
exposed to fewer and weaker wave 
events. Annually, mean wave energy 
and wave power (energy transferred 
across a given area per unit time) are 
highest (approximately 1.3 W/m) be- 
tween November and March and lowest 
(approximately 0.3 W/m) between May 
and September (Figure 2.5). Extreme 
wave events, 1 m or higher, affect shal- 
low water coral reef communities with at 
least an order of magnitude more ener- 
gy than the typical winter waves (Grigg 
et al., 2008). 



' * 1 1 ., „ i + 



in r« 



Figure 2.5. Diagram of Climatological values of wave power (W/m) derived 
from NO A A buoy #51001 located near Nihoa Island from 1981 to 2003. 
Blue circles represent monthly means; blue lines represent wave power 
maxima. Source: NOAA NDBC. 



Significant wave events vary over interannual and decadal time scales. This temporal variability of wave power 
allows expansions and retractions of the spatial and vertical ranges of the same species during relatively qui- 
escent and turbulent years, respectively (Rooney et al., 2008). Over the past 20 years, wave measurements at 
NOAA buoy 51001 (near Nihoa Island in the NWHI) show a pattern of numerous extreme wave events during 
the periods 1 985-1 989 and 1 998-2002 and low numbers of extreme wave events in the early 1 980s and the pe- 
riod 1990-1996. This apparent decadal variability of wave power is possibly related to the PDO (Mantua et al., 
1997). Studies have shown decadal oscillations of various components of NWHI ecosystems (lobsters, monk 
seals, seabirds, etc.) relate to larger scale climate shifts across the North Pacific (Polovina et al., 1995). 



Primary Productivity / Ocean Color 

Productivity in the NWHI is influenced by local and regional factors, and upwelling may occur in response to 
localized wind and bathymetric features (Friedlander et al., 2005). The Monument is located at the northern 
edge of the oligotrophic tropical Pacific, in the North Pacific central gyre ecosystem (Figure 2.6). Regional fac- 
tors are largely influenced by the position of the subtropical front and associated high chlorophyll content of 
waters north of the front (PMNM, 2008). High chlorophyll waters intersect the northern portions of the NWHI 
during southward winter migrations of the subtropical front. The influx of nutrients to the NWHI from these mi- 
grations is considered a significant factor influencing different trophic levels in the NWHI (Polovina etal., 1995). 
It is near the 18°C isotherm, a major ecological transition zone in the northern Pacific. This boundary, also 
known as the "chlorophyll front", varies in position both seasonally and annually, occasionally transgressing the 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Monument boundary and surrounding 
the northern atolls of Kure and Midway 
(PMNM, 2008). 

Movement of this front influences overall 
ocean productivity and resultant recruit- 
ment of certain faunal elements such as 
Hawaiian monk seals and Laysan and 
Black-footed Albatrosses (Polovina et 
al., 1994). The northernmost atolls also 
are occasionally affected by an episodic 
eastward extension of the Western Pa- 
cific warm pool, which can lead to high- 
er summer ocean temperatures at Kure 
than are found in the more "tropical" 
waters of the MHI further south (Hoeke 
et al., 2006). This interplay of oceanog- 
raphy and climate is still not completely 
understood, but is a dynamic not seen 
in most other tropical atoll ecosystems. 
As a result, it provides a useful natural 
laboratory for understanding phenom- 
ena such as periodic coral bleaching and the effects of El Nino and La Nina ocean circulation patterns (see 
ocean temperature). 

Satellite observations of ocean color from the National Aeronautics and Space Administration's (NASA) Sea- 
viewing Wide Field-of-view Sensor (SeaWiFS) reveal a significant chlorophyll front associated with the sub- 
tropical front, with high chlorophyll north of the front and low oligotrophic waters south of the front. These 
observations reveal significant seasonal and interannual migrations of the front northward during the summer 
months and southward during the winter months (Seki et al., 2002). The southward migration of the subtropi- 
cal front generally brings these high chlorophyll waters to intersect the northern portions of the NWHI. During 
some years, these winter migrations of the subtropical front extend southward to include the northern end of 
the NWHI. Additional evidence suggests decadal scale movements in the southward extent of the subtropi- 
cal front. During periods when high chlorophyll waters intersect the NWHI, overall productivity of the affected 
reef ecosystems is expected to be elevated. Changes across many trophic levels of the NWHI ecosystem are 
believed to be associated with these migrations (Polovina et al., 1995). 



Figure 2.6. Diagram of Central Pacific Gyre. The North Pacific, California, 
North Equatorial, and Kuroshio currents along with atmospheric winds gen- 
erate the North Pacific Subtropical Gyre. The subtropical Convergence 
Zone, an area where marine debris is known to accumulate, shifts season- 
ally between 23°N and 37°N latitude. 



OCEAN REMOTE SENSING ANALYSIS: DATA AND METHODS 

This oceanographic assessment is based largely on data acquired from satellite and in situ sensors to char- 
acterize conditions for each of the management areas within the Monument, as well as the larger ecological 
region. For this report, the study area is defined as: north bounding coordinate - 45°N; south bound - 15°N; 
east bound - 145°W; and west bound - 175°E. Spatial patterns in the temperature, temperature fronts and 
chlorophyll were identified, as well as the variability in those patterns. Time series information was also ex- 
tracted from the datasets to investigate trends at a variety of time-scales. 

The Monument includes 10 special management zones and eco-regions. These management regions are 
centered on relatively untouched islands, reefs and atolls that are home to thousands of species, some that 
are found nowhere else on earth. The oldest (approximately 28 million years old) and most northern are Kure 
Atoll, Midway Island and Pearl and Hermes Atoll. These three ecosystems are subject to similar oceanographic 
influences and will be analyzed together as the northern grouping. The adjacent, and next most southeasterly 
grouping - Lisianski Island, Laysan Island, Maro Reef and Gardner Pinnacles - lie along the same latitudi- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



175- 


-180 s -175- -170' -155- -160- -155" -150- 


-145° 


-140" 








r 






I 




^H-40' 


40 -H 












I 


' ^ i" 






1 


-35' 










35 -U 








-30' 


30 B 










I 

25 ~H 








-26- 


-p'-Bj 


m|9 /X 






-20' 




-175" -170" -165' -1 60 -155" 


-150" 


-145° 



Figure 2.7. Locator map of the study area. Cross-hatched regions repre- 
sent the northern, central, and southern analysis areas. 



nal zone and exhibit similar temperature 
and chlorophyll climatological profiles, 
as well as being subjected to similar 
current regimes. These will be analyzed 
together as the central group. The most 
southern and youngest (7-12 million 
years old) of the atolls are French Frig- 
ate Shoals, Mokumanamana Island, 
and Nihoa Island (Figure 2.7). These 
three experience milder SSTs, smaller 
temperature ranges through the year 
and are less impacted by winter storm 
waves. 

The study area encompasses a region 
much larger than the Monument bound- 
aries so as to allow ecosystem-scale in- 
formation to be included in management 
decisions. Much of the oceanographic 
variety in the region of the Monument 
happens well away from the atolls them- 
selves. The northern group of atolls is 
the only which experiences extreme 
events and changes on a fairly regular 
basis. Even though the Monument itself 

is relatively calm compared to the ocean environment around it, the more extreme events of the central pacific 
influence the migration of fauna in and around the Monument atolls, and thus must be considered. 

Data Assembly and Processing 

The majority of remote sensing data products were obtained as monthly composites (mean), and subsequently 
processed into seasonal and interannual monthly means and medians. Seasonal means were calculated us- 
ing the following constraints: winter (January, February, March); spring (April, May, June); summer (July, Au- 
gust, September); and fall (October, November, December). Production of a consistent time series of imagery 
in this manner allowed extraction of data from each source within regions of interest, and for regional averages 
within the bounds of the entire Monument. These time series were extracted for 10 locations in the study re- 
gion (Figure 2.7, orange polygons), as well as the three regional groupings and the Monument as a whole for 
preliminary analyses of episodic, seasonal and interannual patterns. Interannual monthly means were then 
derived from the extracted time series. These data composites over the entire Monument, or for specific loca- 
tions, were created to summarize trends and to highlight episodic and seasonal events useful for interpreting 
biogeographic patterns discussed later in this report. Time series images consist of all monthly mean or me- 
dian data for each month over time. For example, the month of January has an image of mean chlorophyll for 
each of the years 1998 to 2007. These 10 monthly images are then averaged to obtain an interannual mean 
of chlorophyll. Monthly and seasonal anomaly images were produced to illustrate the differences between the 
long-term interannual mean and the monthly image. 

Chlorophyll 

Chlorophyll products were derived from data obtained from SeaWiFS. SeaWiFS data for research and edu- 
cational applications have been available through the NASA since its launch in September of 1997 through 
December 2007. The sensor provides reliable daily observations for the United States at a nominal spatial 
resolution of 1 .1 km for spectral bands encompassing the visible and near-infrared spectrum. The 4 km Global 
Area Coverage (GAC) product was used because data quality and density of the 1.1 km data was insufficient 
in the NWHI region (Figure 2.8). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



The entire SeaWiFS GAC dataset was 
processed with SeaDAS version 5, ap- 
plying the improved algorithms and 
obtaining georeferenced chlorophyll-a 
data at 4-km spatial resolution for the 
central Pacific region. Products were 
created for the central Pacific region 
specifically to include the boundaries 
of the Monument and also encompass 
the surrounding environment to allow 
management and scientific inquiry to in- 
clude the greater ecosystem contextual 
processes. Estimations of chlorophyll-a 
in units of ug/L were derived using the 
standard OC4v4 equation that NASA 
used for global products from SeaWiFS. 
NOAA's Center for Coastal Monitoring 
and Assessment (CCMA) has devel- 
oped algorithms, implemented by NASA 
for standard processing, to improve the 
generation of ocean color data and es- 
timation of chlorophyll from SeaWiFS 
(Stumpf etal., 2003). 



175'- -180- -175- -170- -165 




N6 

I 1 1 1 1 I— < t— H 



Figure 2.8. Example Global Area Coverage (GAC) chlorophyll distribution 
throughout the study region (April 2004). Warm tones represent high chlo- 
rophyll, while cool tones represent lower estimated chlorophyll concentra- 
tions (Units are pg/L). 



Chlorophyll-a is the dominant pigment in 
marine photosynthetic organisms, and 
is referred to simply as chlorophyll within this report. Time series image sets of chlorophyll were created for the 
specified regions in Geotiff format. These time series images are useful for determining trends in algal bloom 
activity and ocean productivity. Final products were projected using the Albers Conical Equal Area (ACEA) 
projection with the World Geodetic System 1984 (WGS-84) datum. The imagery time series generated from 
the SeaWiFS data are monthly medians. Seasonal means were created from the appropriate monthly median 
files resulting in a seasonal image (seasons previously defined), and interannual monthly and seasonal files 
were generated using all monthly images for a particular month (or season) in the time series as input. 

Sea Surface Temperature and Frontal Boundaries 

SST data were developed from the NASA Pathfinder Version 5.0 dataset. This dataset derives a climatological 
grade SST product from Advanced Very High Resolution Radiometer (AVHRR) imagery, which was generated 
from several NOAA Polar-orbiting Environmental Satellites between 1985 and the present. The Pathfinder 
dataset was calibrated for inter-comparison of the temperature data across the entire period, facilitating cli- 
mate and other studies (NASA, 2004; NASA, 2005). Ocean fronts data, regions that delineate the boundary 
between different water masses, were obtained from the Geostationary Operational Environmental Satellite 
(GOES). 

Fronts were generated from GOES 2000-2006 hourly images (averaged to months) sampled to 4 km (Figure 
2.9). Fronts were identified in each image using an algorithm developed by Canny (1986). Front occurrences 
were tallied at each pixel location for a month, season or year as needed. Climatological means were also cre- 
ated. Data products were mapped and subset to the study area. Geotiffs were created using the ACEA projec- 
tion with the WGS-84 datum (Figure 2.9). 



Spatial resolution of the Pathfinder data varies slightly with latitude, with a horizontal resolution of approximate- 
ly 4 km at 35 degrees of latitude. Pathfinder is distributed along a 0.04 degree grid in Cartesian coordinates. 
The original global 4 km data were subset to the study area bounds. Geotiffs were created of the monthly mean 
time series (1 985-2006) using the ACEA projection with the WGS-84 datum. SST image datasets were created 
from monthly means for the region; units are degrees Celsius (°C). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Ocean Currents 

The calculation of SSH is based on a 
reference ellipsoid. This reference ellip- 
soid is a raw approximation of Earth's 
surface, a sphere flattened at the poles. 
This position is determined relative to 
an arbitrary reference surface, an el- 
lipsoid. The satellite altitude above the 
reference ellipsoid (distance S) is avail- 
able to within 3 cm. The SSH, is the 
range at a given instance from the sea 
surface to a reference ellipsoid. Since 
the sea depth is not known accurately 
everywhere, this reference provides ac- 
curate, homogeneous measurements. 
The sea level is simply the difference 
between the satellite height and the alti- 
metric range. 





-180' -175" -170' -165' -160' -155' -150' 


35'- 


■£- - " ■ -' ^- ,^^v?^" ■'''■-' '-■ '■.■ l • '■ 


-35' 


30 -- 

25 - 


^Hf ; -'■ ,L ha 


-30 

-2? 


BBHP^ ' 




1 ' *"fts 




20- 


210 420 B40 Kilometers 

I — i — i — i — 1 — i—i — i — 1 


-20 


-175 ; -170' -165' -160' -155' -150 : 





Figure 2.9. Example average sea surface frontal boundaries for the region 
(April, 2005). Dense clustering of boundaries, and lighter tones indicate 
elevated frontal boundary detection. Note higher frontal activity in the north- 
ern and western reaches of the Monument. 



Merged sea surface height anomaly 
(SSHA) data from altimetry were ob- 
tained from Archiving, Validation and In- 
terpretation of Satellite Data in Oceanography (AVISO) delayed time products of SSH generated from merged 
Topex/Poseidon (T/P), Jason-1/2, ERS-1/2 and ENVISAT missions. Merged SSHA data from altimetry were 
obtained from AVISO delayed time products of SSH generated from merged Topex/Poseidon (T/P), Jason- 
1/2, ERS-1/2 and ENVISAT missions. Weekly (seven-day) and monthly averaged data were used. Monthly 
estimates of vertical SSH from mean sea level were obtained following a simple bin averaging technique. A 
calendar of monthly averaged SSHA data to dissect climatological patterns of space-time variability for the 
region of interest is shown. 



Post processing involved conversion from Network Common Data Format (NetCDF) format to raw binary 
image format, binning of weekly SSHA data, georeferencing and tiff generation of by-products. The binning 
procedure followed a simple arithmetic averaging technique to compute a monthly estimate of vertical SSH 
from mean sea level. Map projection is a 1/4° geographic (lat/long) projection grid, where number of values for 
X (found in file) = 1 ,080 and number of values for Y (found in file) = 720. Final georeferenced products were 
converted to 32 bit Geotiffs (v6). 8 bit Geotiffs were also created for reference. The time period for SSHA data 
analyzed here range from October 1992 to present. 

Winds 

NOAA's National Centers for Environmental Prediction (NCEP) generates global wind data termed "reanalysis 
winds" which are processed using a state of the art analysis system, and are used primarily for long-term cli- 
mate studies. The data were produced at a 2.5 degree spatial resolution. NCEP uses all available atmospheric 
data to model winds every six hours, and is available from January 1, 1948 through the present. Reanalysis 
winds are distributed in NetCDF, and have a time component, latitude, longitude, zonal component (u) and a 
meridional component (v). 

From the u and v wind components, direction (in degrees) and magnitude (in speed; m/s) can easily be de- 
rived. The data are available for 17 different pressure levels. To estimate the winds closest to the atmosphere- 
ocean interface we chose the highest pressure level (1 ,000 millibars). Using the six hour observations, NCEP 
calculates daily, monthly, and annual average and standard deviation reanalysis wind products. NOAA's CCMA 
has created direction and speed images created as 32-bit Geotiffs in geophysical units primarily for qualitative 
assessment purposes and an 8-bit scaled Geotiff version primarily designed for quantitative assessments. 
Monthly, seasonal and annual means for the study region have been created in this fashion. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

The monthly data from NASA's Quick Scatterometer (QuikSCAT) was obtained and processed form from the 
French Institute for Exploring the Sea (IFREMER). Scatterometers, such as QuikSCAT, measure the rough- 
ness of the surface of the ocean, which in turn may be used as a proxy to estimate wind speed and direction. 
Scatterometer data are not available near or adjacent to land. The QuikSCAT NetCDF data files were gener- 
ated from monthly data composites that covered the period ranging from August 1999 though February 2007. 
Seasonal and yearly climatological files were generated through the combination of monthly data composites 
that covered the period January 2000 though December 2006. The data were spatially subset to the study 
area at a spatial resolution of 0.5 degrees per pixel. Each file contained the following parameters: wind stress 
curl (Pa/m), wind speed divergence (m/s), wind speed (m/s), zonal wind speed (m/s), meridional wind speed 
(m/s), wind stress (Pa), meridional wind stress (Pa) and zonal wind stress (Pa; IFREMER, 2002). The result- 
ing data files for each parameter were converted to 8-bit and 32-bit Geotiff format, in the same manner as the 
reanalysis wind products. 



OCEAN REMOTE SENSING ANALYSIS: RESULTS 

Sea Surface Temperature 

Latitude is a primary driver in oceanographic conditions of the Monument and SST is highly correlated with 
latitude. This is evident in the stratification of SST within the Monument moving from north to south. SST analy- 
sis suggests that the 10 management regions can be segregated into three latitudinal subgroups. Mean SST 
highs in August and September are similar for all of the regions, around 27°C, but lows in February and March 
are varied and highlight the latitudinal differences (Figure 2.10). 

The northern atoll group (Kure, Pearl and Hermes and Midway) has a wide range of temperatures throughout 
the year, typically from around 20°C in February to 27°C during the summer; however, temperatures have 
reached as low as 1 6°C and as high as 29°C. This range is one of the widest temperature ranges for any coral 
reef system on the planet (Friedlander et al., 2005). In addition, the northern islands are unexpectedly warmer 
in the summer than the most southern atolls of Nihoa, Mokumanamana and French Frigate Shoals. This north- 
south temperature stratification results in a latitudinal partitioning of flora and fauna causing endemic species 
and species composition changes within the Monument (Polovina et al., 1995; Friedlander et al., 2005). 

Average monthly SST images clearly show the seasonal variation and north-south temperature gradient in the 
NWHI (Figure 2.11). The southern region experiences warmer temperatures compared to those in the north, 
but less variability throughout the year, while the northern atolls have much more variability within and between 
years (Figures 2.4 and 2.11). The southernmost island grouping (French Frigate Shoal, Mokumanamana Is- 
land, and Nihoa island) experience an environment with a restricted range in temperature when compared to 
islands to the north and west; approximately 23°C in the winter to 27°C in the summer. Summer SSTs for this 
group are generally lower than or the same as the northern atoll group. This is possibly due to weaker winds in 
the north, caused by proximity to the North Pacific high pressure ridge, resulting in less mixing with subsurface 
waters (Hoeke et al., 2006). 

Pattern and periodicity in SSTs within the Monument are remarkably stable from 1985-2006 (Figure 2.12). 
Anomalies are rarely more than a degree from the long-term mean and typically occur during the winter months 
(Figure 2.12). Wintertime negative temperature anomalies occurred in June 1987, May 1992, December 1996, 
February 1997 and June 1997, while positive anomalies occurred during the early spring in 1999, 2000 and 
2001 (Figure 2.12). These observed anomalous variations in SST are likely associated with the interactions 
and state of the PDO and ENSO and trend strongly towards the phase of the PDO, either warm or cold. There 
is less observed variability during the summer months; however, prolonged positive anomalies of 1°C during 
the warmest months are likely indicators of coral bleaching (Kenyon et al., 2006a; Kenyon and Brainard, 2006). 
Similar time-series analysis has been performed for data extracts from each of the 10 islands, atolls and spe- 
cial management areas. Results are provided in Appendix I. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




a 



o 

O 



ID 
CO 



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a. 

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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




January 



February 




March 









July 



+ 



August 



1 



September 





November 



+ 



December 



+ 



Figure 2.11. Monthly mean climatological SST, 1985-2006. Color bar is in degrees Celsius. Warm waters encompass the 
entire island chain in summer months but cooler waters intrude on the northern end of the Monument from November 
through May. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



f+ 29 

28 

27 

26 

SST 25 
(degC) 24 

23 
22 
21 
20 



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A A A ./i A 

ii \i_ [it ;|£ y o 



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c_o) c_Q) E. ^ E. 115 E. 115 E. 115 E. 113 E. 113 



c_ c_ 
E. m 



E. m 

£ S & S 6 

N) w W W -^ 



T 3 

S o 



C CD c 

' o 

01 g 05 



Date 



■mean sst 



■grand mean sst 



(deg C) 




o J-, 6 
u. g o) 



-Anomal 



Date 



Figure 2.12. Panel A shows SST derived from NASA Pathfinder AVHRR imagery. Data from the entire Monument have 
been averaged to highlight temporal patterns from 1985 through 2006. Grand mean indicates the "climatological" aver- 
age. Panel B shows temperature anomalies for the same period of record. Green bar indicates the range of one standard 
deviation of the anomaly time series. Peaks that fall above or below this range can be considered departures from "ex- 
pected" anomalies. 

NOAA led Pacific Reef Assessment and Monitoring Program expeditions to the NWHI documented the first 
recorded major bleaching events in the region. The NWHI were impacted by mass coral bleaching during 
late summer 2002 and again in 2004 principally due to a distinct region of higher than normal temperatures 
pervasive in the northern reaches of the Monument (Figure 2.13; Abey et al., 2003; Kenyon et al., 2006). No 
records of mass coral bleaching in the NWHI existed before this time. It was previously thought that the NWHI 
were less susceptible to bleaching due to the high latitude location whereas coral bleaching was documented 
in the MHI back in 1996 (Jokiel and Brown, 2004). During both events, bleaching was most severe at the three 
northern-most atolls (Pearl and Hermes, Midway and Kure), with lesser incidences of bleaching at Lisianski 
Island and farther south in the NWHI. SST data derived from both remotely sensed satellite observations as 
well as in situ buoys from the NOAAs Coral Reef Early Warning System suggest that protracted, elevated SST 
was a likely explanation for the bleaching response. This period of elevated SST coincided with a prolonged 
period of light wind speed, suggesting increased stratification due to decreased wave mixing of the upper 
ocean (Hoeke et al., 2006). 

The time series analysis of SSTs data also suggests that two bleaching events may have occurred undetected 
in the summers of 1987 and 1991 (Figure 2.14). On these occasions, the northern atolls experienced tem- 
perature anomalies of 1°C or greater during the highest mean temperature months of August and September. 
Similar or larger positive anomalies have occurred in other regions and times but likely did not lead to coral 
bleaching because these events did not occur during peak temperature months. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



:■: 



t^n i7j in) .he.' 



,tm -ik- 



i» 






;.- 




SST AnomJv 



I 



Warmer 
+3°C 



-3°C 
Coder 




Figure 2.13. SST Anomaly images for August 2002 (A) and September 2004 (B). These periods of anomalously high 
SSTs in the northern atolls have been associated with coral bleaching. Source: NOAA GOES Imager. 








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Q)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)CQ)C 



°? rV, 00 i, Oq A, TO 



Ol "' O) 



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CD 

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~ JJ ^. w ,i. UJ ,i. JJ ^. JJ ,„ UJ :. UJ ;. n; ,„ UJ : n UJ ,\- IU .a u^ f \^ i^j f \^ 

PPcn°3CT) 3^ 3 m pD CD cp o cD^cp ls jCp a3 cpi^cp C j 1 cp cr ]Cp^,cp 



K> '~ CO 



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■ Midway 



Ol "' 05 



Date 



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N> « CO 



K> '~ CO 



oi " en 



oi "' 01 



Figure 2. 14. SST Anomaly images for September 1987 (A) and September 1991 (B). Distinct SST peaks in the northern 
atolls can be seen in the time series data (C). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Sea Surface Temperature Fronts 

Bottom topography, ocean current confluence, variable wind stress and heat-water exchange across the sea 
surface produces patterns of vertical circulation, fronts or eddy-like motions that can affect biological distribu- 
tions (Kazmin and Rienecker, 1996; Polovina, et al., 2001; Nakamura and Kazmin, 2003). Fronts are created 
by a variety of physical processes and have a wide range of biological consequences. Afrontal system denotes 
areas of water mass convergence and usually produces zones of downwelling and upwelling flow. These verti- 
cal displacements have considerable ecological effects because environmental gradients, such as light, pres- 
sure, temperature, salinity, oxygen, nutrients, etc., are steepest in the vertical axis of the water column. 

Vertical motion in fronts is often highly localized and can be easily identified with remote sensing techniques 
(Ullman and Comillon, 1999). Areas with frontal activity tend to be areas of high biological activity (Polovina et 
al., 2001; Nakamura and Kazmin, 2003). The vigorous mixing of the water column at water mass confluence 
stimulates phytoplankton photosynthesis and sustains concentrations along the frontal zone (Savidge, 1976; 
Savidge and Foster, 1978). In response, zooplankton tend to concentrate along the fronts and are preyed upon 
by higher trophic groups. Marine birds, marine mammals and fish often aggregate at frontal areas as well. 
Tracking and catch studies have shown that key apex predators, swordfish, albacore tuna and loggerhead 
turtles, use these fronts for forage habitat on long distance migrations through the North Pacific (Bigelow et 
al., 1999; Laurs et al., 1984; Polovina, et al., 2001). Additionally, it has been shown that biological diversity is 
positively correlated with thermal fronts (Kazmin and Rienecker, 1996) 

SST fronts are an important marine ecological feature for the overall eco-region of the Monument and the 
dynamic environment created by the fronts support many species that reside near or in the Monument. Large, 
persistent front generating regions are important for species transiting through the Monument during migra- 
tion. However, front concentration within the Monument is relatively low, reaching highest levels in spring and 
primarily in the northern end of the Monument where the subarctic and subtropical gyres interact. Consistent 
with SST observations, SST frontal data indicate a seasonal patterned and latitudinal division within the Monu- 
ment. Figure 2.15 illustrates the north-south movement of frontal concentration throughout the year. The north- 
ern region of the Monument experiences an active frontal season from December through April, usually with 
a peak in March. Even during the peak frontal probability period the southern region experiences few fronts. 
In mid-summer the frontal activity zone retreats far enough north that even the most northern atolls may not 
experience any significant fronts. 




Figure 2. 15. Average monthly sea surface frontal boundaries for the region. Dense clustering of boundaries, and lighter 
tones indicate elevated frontal boundary detection. Note higher frontal activity in the northern and western reaches of the 
Monument. Color strip denotes calculated frontal probability. 

These pelagic habitats are poorly understood and warrant additional studies to improve the knowledge base 
for ecosystem management. It is possible with remotely sensed data to track these frontal zones over large, 
difficult to access regions through time. The interaction of these productive zones with the biota of the Monu- 
ment may hold clues to long term sustainability of the Monument reef systems. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Chlorophyll 

Chlorophyll concentrations are relatively low throughout the Monument, exhibiting oligotrophic characteristics 
common in open ocean environments. Even at these low levels, seasonal and latitudinal patterns are evident 
in both the chlorophyll images and the time series charts. The images shown in Figure 2.16 illustrates the tem- 
poral distribution of chlorophyll. This distribution is determined by the location of the convergence zones of the 
subarctic and subtropical gyres. A major area of productivity associated with this convergence area, referred 
to as the transition zone chlorophyll front (TZCF), contributes significantly to the variable productivity of the 
Monument region. 



0.18 i 



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0.06 - 




0.04 



Jan 



Feb 



Mar 



Apr 



May 



Jun 



Jul 



Aug 



Sep 



Oct 



Nov 



Dec 



Figure 2. 16. Average monthly chlorophyll concentration (mg/m 3 ) values for the study region. 

The TZCF is a high productivity zone in the open ocean used by many species as an important feeding and mi- 
gration zone. The TZCF has been identified as a chlorophyll concentration of approximately 0.2 ug/L or greater 
as measured from satellite (Polovina, 2005). The TZCF only rarely transits far enough south during its winter 
southerly shift to interact directly with the northern regions of the Monument. Even in these instances chloro- 
phyll concentrations within the Monument generally do not reach as high as 0.2 ug/L However, both perma- 
nent and seasonal residents of the Monument do make use of this productive zone and more southern tracks 
of the TZCF have been correlated with higher fish catches in the Hawaiian Islands (Polovina et al., 2001). 
Coral reefs are also impacted by these occasional pulses of higher chlorophyll through population increases of 
the coral eating sea star, Acanthaster planci commonly known as the crown-of-thorns sea star (Hoeke et al., 
2006). These periodic increases in productivity within the Monument likely have meaningful consequences to 
management of the atolls and reefs of the Monument. 



Mean chlorophyll concentrations have cyclical temporal variability. Higher chlorophyll concentrations move 
south into the Monument boundaries during the winter months and retreat far north in the summer. The north- 
ern region experiences the largest variability in mean concentration from high season to low season ranging 
from highs of nearly 0.18 ug/L to lows below 0.06 ug/L. The central and southern regions exhibit a more con- 
stant environment with chlorophyll concentrations ranging between 0.07 and 0.11 ug/L. In addition to seasonal 
changes, the northern regions also exhibit high interannual variability in chlorophyll concentrations. The north- 
ern atolls experienced very low concentrations of chlorophyll in 1999, 2000 and 2001, but also relatively high 
concentration years in 1998, 2003, 2004 and 2005 (Figure 2.17). Note that even the highest mean monthly 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

chlorophyll concentration is below the threshold of 0.2 |jg/L used to identify the biologically active TZCF. Time- 
series analysis has been performed for chlorophyll data extracts from each of the analysis regions, and results 
are provided in Appendix II. 

While these values are low relative to coastal continental chlorophyll concentrations, these chlorophyll blooms 
offer sustenance to support a diverse community around the atolls of the Monument. The southern atoll group 
exhibits little variance through time and can be characterized as having generally low chlorophyll concentra- 
tions, usually between 0.06 and 0.09 ug/L. The central region exhibits slightly higher variability and a broader 
range in chlorophyll concentrations when compared to the south, likely owing to its position as a transition zone 
between the subarctic north and the subtropical south. 



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Figure 2. 1 7. Time-series for chlorophyll (top) and chlorophyll (mg/m 3 ) anomaly (bottom) in each analysis region of the 
Papahanaumokuakea Marine National Monument (1997-2006). Source: SeaWiFS. 



Trends in chlorophyll concentration anomalies can be correlated with the PDO and ENSO via the multivariate 
ENSO index (MEI). Chlorophyll concentrates in the more northern atolls are positively correlated with the PDO 
index (p=+ 0.404) where as the southern atolls are negatively correlated (p=-0.02). The rank correlation com- 
pares the relative size and direction of the indicators. For example, in the northern regions from 1998-2002, 
relatively strong cool phase PDO index values (negative) are positively correlated with periods of relatively low 
chlorophyll concentration anomalies (Figure 2.18). This correlation is expected as a cool phase PDO is typical- 
ly indicated by higher SSTs in the north central Pacific (Mantua et al., 1997), and warmer SSTs are associated 
with lower levels of chlorophyll production. An inverse relationship between chlorophyll concentrations and the 
PDO index in the southern atolls is explained by the typically cooler tropical waters in the eastern Pacific during 
cool phase PDO and the association of higher chlorophyll counts and cooler waters. Additionally, PDO effects 
are primarily seen in the more northern latitudes (Mantua et al., 1997), so an increasing strong relationship with 
increasing latitude is not entirely surprising. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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Figure 2.18. Plots of rank-ordered chlorophyll anomaly and PDO Index 1997-2007 values for the northern (top), central 
(middle) and southern (bottom) regions of the Monument. Lines represent non-linear trends through the time-series to 
highlight the degree and nature of correspondence. Poly is a 3rd order polynomial fit through each time-series (trend). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Six El Nino events occurred during the time series analyzed here, including: 1987, late 1991- early 1992, late 
1997-early 1998, late 2002-early 2003, late 2004-early 2005 and late 2006 (National Weather Service, 2007). 
The 1997-1998 was possibly the most intense in the 20th century; however, the northern region of the NWHI 
were more strongly influenced by the weaker events of 2003, 2004 and 2005. This is possibly due to interac- 
tions between PDO and ENSO. The El Nino events are evident in the SST time series throughout the Monu- 
ment but are particularly apparent in the chlorophyll time series of the northern atolls. The 1997-1998, 2003, 
2004 and 2005 El Nifios led to much higher chlorophyll concentrations than normal in the northern half of the 
Monument. During El Nino events, the Aleutian Low pressure system is more intense and extends south into 
the Monument region, resulting in stronger winds, more mixing and cooler SSTs (Bromirski et al., 2005). La 
Nina phases of ENSO show opposite characteristics in the Monument region leading to warmer waters and 
lower chlorophyll concentrations. La Nina events occurred in 1988, early 1989, and late 1998-2000. The La 
Nina events produced slight decreases in chlorophyll, but not of the magnitude of increase seen in the El Nino 
years. 

Wind 

Winds of the NWHI are generally dominated by the Trades, a persistent system that blows from the northeast 
to the southwest. These winds move from the Americas to Asia between the equator and 30oN, and are re- 
markably consistent throughout the year. Meridional variability is relatively weak and is dominated by Coriolis 
Forces which set up the North Pacific Gyre. Meridional winds are defined as the directional component along 
the local meridian, and are positive if from the south, and negative if from the north. Likewise, the zonal wind 
component is positive if it blows from the west and negative if from the east (i.e., Westerlies). An analysis of 
wind data (1985-present) revealed strong NWHI zonal variability in the NWHI, and three distinct components 
in the region. 

The southerly portion of all monthly wind climatologies - from 15°N to approximately 30°N - is dominated by 
the trade winds which are associated with warmer air and the Pacific High Pressure System (Figure 2.19). 
North of the trade wind zone is a transitional area extending from approximately 30°N to 40°N of weak variable 
winds known as the North Pacific Doldrums, which ancient mariners referred to as the "Horse Latitudes". In the 
northern portion of the study area (above 40°N), winds prevail out of the west southwest, which is associated 
with cooler air and dominated by the Aleutian Low pressure system. The location of the three respective areas 
migrates north and south seasonally, and is a major forcing function of regional oceanographic conditions. In 
addition to the analysis of monthly winds described above, average annual winds were calculated and ana- 
lyzed for the study area. Most years exhibited only modest changes in interannual variability. The one notable 
exception was the intense El Nino event of 1 997. Figure 2.20 shows the annual mean wind vectors from 1 996, 
1997 and 1998, sequentially. The Trade Winds in the annual mean of 1997 a near full directional reversal. In 
1 996 and 1 998 the Trade Winds were out of the east northeast, as expected. In 1 997; however, the winds were 
strongly blowing out of the south southwest. Also notable is that the general circular pattern of the winds, which 
typically form a gyre in the North Pacific, were absent. Throughout the analyzed region winds were out of the 
southwest which is a drastic difference from all other calendar years analyzed (1 985-2007). 1 997 also showed 
remarkable zonal variability along 30°N. West of -170° the magnitude of the winds were much less than they 
were to the east. 

Circulation around the high pressure is clockwise in the northern hemisphere and counterclockwise in the 
southern hemisphere. The high pressure in the center is due to the westerly winds on the northern side of the 
gyre and easterly trade winds on the southern side of the gyre. These cause frictional surface currents towards 
the latitude at the center of the gyre. The buildup of water in the center of the gyre creates equatorward flow in 
the upper 1 ,000 to 2,000 m of the ocean, through rather complex dynamics. This equatorward flow is returned 
poleward in an intensified western boundary current. The North Pacific Gyre comprises most of the northern 
Pacific Ocean, and occupies an area of approximately 34 million km 2 . The Gyre has a clockwise circular pat- 
tern and comprises four prevailing ocean currents: the North Pacific Current to the north, the California Current 
to the east, the North Equatorial Current to the south, and the Kuroshio Current to the west. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




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winds. Wind field data are superimposed on a false-color image of average monthly SSTs (warm tones=warm water, cool 
tones=cool water). Arrow size denotes relative wind strength. Source: AVHRR climatology. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




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event from 1997 shows relatively drastic variability from the preceding and subsequent years. Arrow size denotes relative 
wind strength. 



Sea Surface Height and Currents 

The height of the sea surface is determined by the mass of water at a given location and by the water's density 
(a function of temperature, salinity and pressure). Space based altimeters such as the Jason and Topex/Po- 
seidon missions measures changes in SSH due to both of these factors - redistribution of mass and changes 
in density. On seasonal to interannual time-scales, density changes are the largest contributor to sea level 
variability. In the tropics they are the dominant one (Gilson et al., 1998). 

Ocean currents can increase SSH by up to a meter higher over the surrounding area. Currents can therefore 
be mapped by measuring height variations. A view of the global ocean circulation shows currents circulating 
around elevations and depressions in SSH. Currents flows around positive SSH in a clockwise direction in the 
Northern Hemisphere, and in a counterclockwise direction around negative SSH (the opposite occurs in the 
Southern Hemisphere). Figure 2.21 shows an example AVISO derived SSHA image for the study region during 
December 2005. Warm tones indicate regions of surface height elevation, while cool tones highlight surface 
depressions. Current vectors derived from SSHA are superimposed on the image to show modeled surface 
currents (geostrophic flow). As described, note the counterclockwise rotation around the blue tones. 

Numerous studies have summarized the positive association between SSH maximums and SST maximum in 
open ocean environments (Jones et al., 1998; Wilson and Coles, 2005; Fu 2004). Given the setting of the Mon- 
ument in the middle of the Pacific Ocean, it is therefore not surprising that SSHA patterns correlate with SST 
seasonal patterns. The difference between the measurements is that SSH are more a measure of the heat 
(energy) that is stored in the ocean below while SST is a surface measurement that tells us about interactions 
with the atmosphere on a more immediate time scale (JPL). The ocean energy below the surface reflected in 
the SSHA influences surface events over much longer periods and areas. 

Localized and episodic SSHA events can be attributed more readily to wind events (Di Lorenzo et al. 2008; 
http://www.ig.utexas.edu/research/projects/od_sst/) but larger regional SSHA patterns are clearly tied to SST. 
The direct influence of SSHA on the atolls of the Monument can be found in generation of winds and a current 
related to transient local eddies and fronts (Seki et al., 2002). Mesoscale SSHAs can be used to identify areas 
of convergence that can indicate upwelling and productive ocean zones concentrating and attracting species 
throughout the food web (Seki et al., 2002). These mesoscale features are embedded in larger scale frontal 
zones that scan large quadrants of the Pacific Ocean and change over periods of months and years. The PDO 
is a long period ocean feature reflected in SSHA. The PDO waxes and wanes between cool and warm phases 
approximately every five to 20 years. In the cool phase, higher than normal SSH caused by warm water form a 
horseshoe pattern that connects the north, west and southern Pacific, with cool water in the middle (JPL). The 
NWHI lie along the typical boundary between cool and warm waters during a cool phase PDO. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

SSHAs in the Monument do have a seasonal signal with higher anomalies occurring in late summer and lows 
in the spring (Figure 2.22). The regional differences observed in the other oceanographic factors are not as 
pronounced with SSHA. The northern regions do experience a less pronounced swing in anomalies from high 
season to low season. 




Figure 2.21. Example sea surface height anomaly image and associated surface currents in the study region (December 
2005). Red tones indicate "peaks", while blue tones indicate "valleys", and arrows indicate direction of flow. Muted poly- 
gons delineate Monument "Special Protection Areas". 







onument 








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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



EXISTING DATA GAPS 



Overall there is a need for basic information on spatial and temporal patterns of water movement, quality and 
characteristics within the NWHI at a range of scales. Moving forward it is important for resource managers to 
have a unified hydrodynamic model to describe connectivity, identify seasonal areas of oceanographic pro- 
ductivity, detect change in the pattern and scales of movement, dispersal and recruitment of living resources 
at various life stages, identify and document variability in larval and nutrients sources, understand debris dis- 
persal and establish management units within the NWHI. Specific opportunities include research to improve 
the understanding of: 

• carbon, nitrogen and phosphorus in the ecosystem and the transfer to higher trophic levels; 

• community changes that will result from alterations to reef structure by major ocean/atmosphere events; 

• discerning anthropogenic impacts from natural variability of the physical ocean environment; 

• PDO/ENSO events and effects; 

• geomorphological and sedimentalogical processes affecting reefs and terrestrial areas; 

• dispersion patterns of key pollutants; and 

• physical and biological effects of extreme events on the ecosystem. 



CONCLUSIONS 

The Monument is a unique open ocean ecological observatory, relatively free from the activities of humans 
and so large it encompass multiple overlapping and interacting marine ecosystems. The Monument's position 
near the shifting boundary of the oligotrophic North Pacific Central Gyre and the productive waters of the North 
Pacific Subpolar Gyre makes it an ecosystem that is influenced directly by climate systems that vary greatly in 
time, over years and decades, and space, over hundreds of kilometers. The atolls and islands are subject to 
typical yearly seasonal patterns as well as the larger climactic cycles of the PDO and the ENSO. 

The ecosystems of the Monument are linked by these circulation patterns but are also stratified by their dis- 
tance from the frontal zone where the gyres meet. There are three latitudinal groups within the Monument 
bounds. The northern group, Kure, Midway and Pearl and Hermes; the central group, Lisianski, Laysan, Maro 
Reef and Gardener Pinnacles; and the southern group, French Frigate Shoals, Mokumanamana and Nihoa. 
These groups exhibit similarities in all the factors that were examined. The environmental factors examined in 
this report are all directly impacted by the changes in the basin wide circulation system. These linkages happen 
over monthly, seasonal, annual and longer term climactic time intervals. Examination of chlorophyll-a, SST, 
SST fronts, and SSHA over climatological and monthly periods make it clear that these factors are linked. 

In addition, these factors correlate with patterns and changes in climactic scale events such as the PDO and 
ENSO. These large-scale oceanographic forcing mechanisms change the characteristics of water temperature 
and productivity across the Pacific, and have a significant effect on the habitat range and movements of pelag- 
ic species in the NWHI. Tuna are often concentrated near islands and seamounts that create oceanographic 
divergence and convergence zones, which in turn tend to concentrate forage species (PMNM, 2008). Sword- 
fish and numerous other pelagic species tend to concentrate along food-rich temperature fronts between cold 
upwelled water and warmer oceanic water masses (Polovina et al., 2001 ). These frontal zones also have been 
determined to function as migratory pathways for loggerhead sea turtles (Polovina et al., 2001, 2004). 

Hundreds of thousands of seabirds breed in the Monument and are primarily pelagic feeders. The fish and 
squid they consume are generally associated with schools of larger predatory fish described above. While both 
the predatory fish and the birds are capable of foraging throughout their pelagic ranges (which encompass 
the entire Monument and tropical Pacific Ocean), the birds are most successful at feeding their young when 
they can find schools of predatory fish within easy commuting range of the breeding colonies (Ashmole, 1 963; 
Feare, 1976; Flint, 1991). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Recent analyses of SeaWiFS remotely-sensed ocean color data shows an expansion of low productivity ocean 
water worldwide (Polovina et al., 2008). This expansion of low chlorophyll water has reached the Hawaiian 
Archipelago and has implications for the productivity of the entire ecosystem. These oligotrophic areas are 
expected to continue to expand with future global warming forcing (Polovina et al., 2008). 

While there is well documented small-scale (i.e., local) variation in population structures (Ashmole and Ash- 
mole, 1967; Boehlert, 1993; Johannes, 1981), the large-scale patterns in oceanography described in this 
chapter provide a dominant force in modulating these populations. Finer-scale, stochastic processes also op- 
erate within this climate driven construct. It is important to note that these processes cannot easily be resolved 
using remote sensing technologies (e.g., wind wakes, wake eddies, etc.), yet are what likely produce pelagic 
"habitat" conditions that attract individuals and sub-populations to a given region, island, atoll or pinnacle. The 
following chapters featured in this document will provide a more detailed view of the abundance, distribution 
and temporal "behavior" of biological populations inhabiting the NWHI. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 
APPENDIX I. SEA SURFACE TEMPERATURE (AND RELATED) TIME SERIES PLOTS 



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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 





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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Monthly Mean Sea Surface Temperature for Pearl and Hermes 1985-2006 



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m 


r 


m 


r 


II) 


r 


m 


-1 


i 


-1 


i 


-1 


i 


-i 


o 


o 


o 


CJ 


o 


o 


o 


— * 




NJ 




OJ 




-^ 



C ID C ID C 



O 
05 



-AnomsHy 



Date 



Seasonal Mean Sea Surface Temperature for Pearl and Hermes 1985-2006 



01 
■D 



(0 
(0 



28.0 




•meansst grand mean sst 



Date 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Monthly Mean Sea Surface Temperature for Lisianski Island 1985-2006 




«SgSs3S3gggSg§s2gSgSgSgggS»5gSggg§22oS&Sgggggg 



■mean sst grand mean sst 



Date 



Monthly Mean Sea Surface Temperature Anomalies, Lisianski Island 1985-2006 




-Anomaly 



Date 



Seasonal Mean Sea Surface Temperature for Lisianski 1985-2006 




scoscoswswscos- 
3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 
£g£g£g£g£g£g£g£g£g£g£g£g£g£g£go 

gggS^ggggg^gggggggggg^gggggg 



w s w s w ■> _. 

" = 3 



w s w s w 



r-o 



M ., N> ., NJ 
O N3 M 

O g O g O X ._ 



.='3 = 3 

NJ ., NJ ., NJ ., N> ., 

ogogogog 

"g^g^g^g 



■mean sst grand mean sst 



Date 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Monthly Mean Sea Surface Temperature for Laysan Island 1985-2006 



29.0 




3T ZST ZST 3T 3T 3T 3T 3T 3T 3T 3T 3T 3T 3T 3T 3T 3T ZST ZST 3T 3T 3T~ 
• 00 ' 00 ' 00 ' 00 ' 00 1 CO 1 CO ,'„ CD ,'„ CD ,'„ CD ,'„ CO ,'„ CO ' CD ' CD ' CD ' O ' O ' O ' O ' O ' O ' O 



00 £ CD g CD »J CD 
CD * O U -1 NJ 



<D CD ° O ° - ° 
CO o -^ K1 



Date 



■meansst grand mean sst 



Monthly Mean Sea Surface Temperature Anomalies, Laysan Island 1985-2006 



o 
2_ 

l- 

C/5 
C/5 




Date 



Seasonal Mean Sea Surface Temperature for Laysan Island 1985-2006 



28.0 




ScScScScScScScScScScScScScSc 

= 3 = 35.3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 = 3 

i^i^cBc^isisic^imisisisisi^ic^i^ 

Date 



oocoScoScooocoScdScd 

uioogooSSoooooogoogcD 

ai u 'o)^^i oo^cd^o 

■meansst grand mean sst 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Monthly Mean Sea Surface Temperature for Maro Reef 1 985-2006 



28.0 
27.0 

o 26 -° 

S? 25.0 

■o 

H 24.0 

(0 

w 23.0 

22.0 

21.0 

20.0 


















































































































































































































































































































































































, ] 


































































DCS 
DTD 

,'~ to ■ 



CO ^ CD 



CD 



C 0) 
T 3 
O ' O 



O 2 O 



C CO c 

— 3 — 
O ' o 



■mean sst grand mean sst 



Date 



Monthly Mean Sea Surface Temperature Anomalies, Main® Reef 1985-2006 




-Anomaly 



Date 



Seasonal Mean Sea Surface Temperature for Maro Reef 1985-2006 



O) 
CI 



CO 

CO 




1_ -: C 

^ 3 = " 



w ?? g g? 3 



-mean ssf grand mean ssf 



d-'d-'d-'d-'d-' 

. - 1 . - 1 . M r^ N> r^ M 

Date 



3 = 3 
ogogogogogo 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Monthly Mean Sea Surface Temperature for Gardner Pinnacles 1985-2006 



29.0 



28.0 



27.0 



O 26.0 

01 

o 

a 25.0 



% 24.0 \- 
23.0 



22.0 - 



21.0 



vl y y if u \i % y i _J_ J_ % " 1 " 



CO 



CO 



CO 



CO 



CO 



CD 



CD 



CD 



CD 



CD 



CD 



CD 



CD 



CD 



CD 



Sffi2fflSs8S8i6S6«Ssio(qS«5i8(iiffl$«^®ffl«So o J O lo o u c)^o ,g | 



en 



o w ^ 



en 



Date 



en 



■mean sst grand mean sst 



Monthly Mean Sea Surface Temperature Anomalies, Gardner Pinnacles 1985-2006 




CO 
-J 



CO i CO 
CO g CD 



-Anomaly 



m c__ 

Date 



gs 



c 


c 


c 


c 


c 


c 


m 


r 


II) 


c 


id 


r 


-i 


i 


-l 


i 


-i 


i 


o 


o 


O 

cn 


cn 


o 


o 



Seasonal Mean Sea Surface Temperature for Gardner Pinnacles 1985-2006 



28.0 




w s w s w ■> CO 

c S c £ c £ c 

3 = 3 = 3 = " 

CD 



3 

ffiSioSioCoioSiD 

oogooS3oooooogooS_ 



_£cSc£c£c- 

3 = 3 = 3 = 3 = 3 = 3 = 3 

'gcSgco'icD'icD'icD'icD'gcD'gcD'gcD'gS 



3 = 

'CD^CD^CD^O^ 

cd">cd£cd£o° 

-^°°C» (D CD 



N) ., N) ., M ., N> ., N) ., N) ., 

o2o°o°o°o°o° 



•mean sst grand mean sst 



Date 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Mean Sea Surface Temperature for French Frigate Shoals NMS 1985-2006 



Ol 

■a 



CO 




01 C 01 C 01 
1 00 1 00 A, 

8 « § °> 5 



C ED 
T 3 

^| 00 
00 



c oi c oi 

T 3 T 3 

CD I CD ' 

00 § CO § 






c ED 
T 3 

2 tb 



nana 

•7- -1 -r- -[ 



£g 



Sg 



E m 
g to 

on 



-mean ssf grand mean ssf 



c_c_c_c_c_c_c_c_ 
C01C01C01C01 

Date 






1 t I 

- E m E 



Mean Sea Surface Temperature Anomalies, French Frigate Shoals 1985-2006 




0) 



_ c o> c oi 

00 g 05 g 03 

01 w 01 u ' -~l 



c o> c oi 

23 1 « i 



C 03 

T" "I 



§g 



E m 
I g 



C 01 C 0) C 0) c 



to 



ss 



0) 

CO g i 

CO w *. 



0) c 

3 7" 
CO .L CO 



01 C 0) 
3—3 

cb S tb 

Ol u ' ~-l 



C 0) C 01 



C 0) C 0) 

S|||tb|§Sc 



c 01 

— 3 



CQ1C01C01CQ1 



01 c: 
3 T 



6g6g6g6°6g 



-Anomaly 



Date 



Seasonal Mean Sea Surface Temperature for French Frigate Shoals 1985-2006 



28.0 




ScSc£c£c£c£c£c£c£cScSc£c£c£c£c£c£c£cScSc£c£c 
g£g£g£££g£g£g£g£g£g£g£g£g£g£g£gogogogogo^ 



cn 00 
01 



01 



00 00 00 

CD 



CO 



CO ^ CO 

o 



M 



CO 



CO 



cn 



Ol 



CO 



CO 



CO 



0^0 

O -1 



w 



Nl 



CO 



CO 



01 



01 



Ol 



Ol 



•meansst grand mean sst 



Date 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Monthly Mean Sea Surface Temperature for Mokumanamana Island 1985-2006 



CI 



(0 

CO 



.0 
.0 
.0 
.0 
.0 
.0 
.0 

n 
























































































































































































































tl \ I Ifl 1 1 \ 1 1W 



















































































































































































































1) C 11 C ill 

^ — 1 — 1 

00 en °° m °° 

01 w (3) ™ S 



CttlcaiCttlCtt) 

00 ' 00 i 00 ' CD ' CD 
^googcDgo^- 



E M 



-mean sst grand mean ssf 



caictticttictticaictti 

cb^cb^ltcbgcbgcb^cbg 



3 T 



c_ c_ c_ c_ 

C ttl C ttl 



SB 

CD t O 

co ^ o 



3 T 



1 C II 

3 T 3 

O ° O 

NO M CO 



c_ c_ c_ c_ 

c ai c ai 



C ttl c 

T 3 T 

' O 

01 ° o> 



Date 



Monthly Mean Sea Surface Temperature Anomalies, Mokumanamana Island 1985-2006 




c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_c_ 

cflicflicflicaicaicaiccucaicaicaicaicflicflicflic 
"T3"r3T3"r3"r3"r3"r:3"r3"r3"r3"r3"r3"r3"r3T 
CO ' CO ' CO i, CO ' CO ' CD ' CD ,'„ CD ,L CD ,'„ CD ,'„ CD ,L CD ,L CD ,L CD '- CD 



01 



01 



CO 



CO 



CD 



§ 5 2 = J8 » 



CO 



CD 



01 



01 



CD 



CO 



CD 



ttl 


c 


ttl C ttl 


c 


(i> 


c 


(11 


c 


ttl C ttl 


r 


-1 


1 


3 "T 3 


i 


-i 


i 


-1 


i 


=1 T =1 


i 


o 
o 


o 
o 


6 2 6 


o 


o 

CO 


o 

CO 


o 

5 


o 

5 


6 S o 
en " oi 


o 

01 



-Anomaly 



Date 



Seasonal Mean Sea Surface Temperature for Mokumanamana Island 1985-2006 





27.0 
26.0 
25.0 






























































































A 




A 
















A 
















ft 




ft 






ft 




















ii 








o 

CI 
CI 

2, 




' 




/ 




J 


l\\\\l 


' I 




' n 


/ 






' r 


H 
CO 
CO 


24.0 






23.0 
22.0 



















































































































































































SW$W$WSW<WSWSWSWSW$W$WSW$WSWS<f)SWSWS<f)S(f)SWSWSW 



CD -i 
00 CD 



_k (O -i 

CD 00 CD 



cnoo£°°^°°°°°°S°°5cD 



CD 
CO 



CD 



- 1 . M 



CD 
CO 



O 
CO 



■mean sst grand mean sst 



Date 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



CI 
■D 



V) 
V) 



29.0 
28.0 
27.0 
26.0 
25.0 
24.0 
23.0 
22.0 
21.0 



Monthly Mean Sea Surface Temperature for Nihoa 1985-2006 






c so c a 



m 

i 00 i 00 J. 

8 " S °> ™ 



I I 

c m 

T 3 

00 i, 



c_c_c_c_c_c_c_c_c_c_c_c_c_c_ 

CD)CD)CD)CD)CD)C£DC£D 



0° ™ 



oo a. cb 
co JS o 



E m 



<b 2 



ss 



Sg 



g cb 
01 



■meansst grand mean sst 



c_ c_ 

C 01 

T 3 . 

CD ' CD ' CD 

Date 



C_C_C_C_C_C_C_C_C_C__ 

ca>ca>ca>ca>ca>ca> 



c cu c a) 

— 3 T 3 

=3 S § ' 



i i i 

C CD C 
— 3 T 



Monthly Mean Sea Surface Temperature Anomalies, Nihoa 1985-2006 




c to c to 



-Anomaly 



C 0! 



C 01 
— 3 



. C B " 

3 — -i 



COJCCUCQJCQJCQ) 
T--5 T - -! T--5 T--5 T - ~1 



C 01 - 



01 

g cb 
^ 01 



8J 



C 01 



r 


(11 


r 


to 


r 


(1) 


C 


(1) 


r 


to 


T~ 


-1 


i 


-1 


1 


-1 


i 


-1 


i 


-1 


CD 

s 


CD 
CO 


CD 
CO 


CD 
CD 


CD 
CD 


O 
O 


o 

o 


o 


o 


o 



C CD C 
7" 3 7" 

O A, o 



to 


r 


to 


c 


(11 


r 


-1 


i 


-1 


i 


-1 


i 


o 


CJ 


o 
en 


CJ 
01 


o 

01 


o 

01 



Date 



28.0 



27.0 



5J 26.0 

o 

E 25.0 

H 
W 
« 24.0 



23.0 



22.0 



Seasonal Mean Sea Surface Temperature for Nihoa 1985-2006 




£c£c£c£c£c£c 

= 3 = 3 = " 

CD 

CO 



c£c£c£c£c£c£c£c£c£c 
333333333 = 3 = 3333333 

^CD^CD^oKoKoKoKoKoKoK 
(D CD <D CD (D 
CDgcDgcDgoSogogogogogo 



3 = 3 = 

CO ^ „ J io - 1 <" 
00 CD 2 CD £ CD ~ 



3 = 



.. <■ CO S W S C/3 

c £ c £ c £ c 

3 = 3 



(fl ?(» JO) 
- £ c £ c: 



l_ _: c - 

3 = 3 = 



en 



CD ™ CD CO CD 

co rj co co 00 

O) ^ -si 00 



>_ _: C £ C - 

_ 3 = 3 = 3 = 

OOOogggggcDgcDScDgcDSggg^^ggg 



■mean sst grand mean sst 



Date 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



APPENDIX II. CHLOROPHYLL (AND RELATED) TIME SERIES PLOTS 

Monthly Median Chlorophyll for Kure 1997-2007 



(191;. 
























J 














— ' n 9n - 
























/ 












I 


























* l\« 


_____ 

-5- 




A. 






j 


U1S 
















_ A 


< it 


fl 




J 


0.05 — 








i i ¥ i 


i i i i 






' 






i i 


I I I I I 


I I I I I I I I I I I I I 





n 


c_ 


> 


c_ 


O 




c_ 


T> 


C_ 







C_ 


> 


c_ 




n 


c_ 


> 


C_ 



n 


c_ 


> 


c_ 


O 

n 


c_ 


f> 


c_ 




n 


c_ 


> 


C_ 







c_ 


> 


c_ 




n 


c_ 


> 


c_ 




n 


c_ 


> 


c_ 


0) 


■d 


C 


m 


T. 


C 


m 


■0 


c 


0) 


■D 


c 


0) 


■D 


c 


U) 


- 


c 


01 


- 


c 


0) 


-0 


c 


0) 


■D 


c 


ai 


■0 


c 








1 








1 
























1 








1 








1 
























■ 


(.0 


(0 


CD 


_ 

co 


CO 


(O 


CD 


CD 
CD 


CD 








CJ 




O 


O 


O 











O 



M 


O 


n 






co 


O 





O 








n 





CJ 
01 


O 


O 


O 



03 


O 


!*) 








N 


co 


CO 


CO 


CD 


CD 


CD 





CJ 





_ 


_i 




_i 


K> 


ro 


ro 


co 


CO 


CO 


*■ 


-1^ 


-t. 


en 


en 


en 


(3) 


0) 


O) 


N 


-~j 








































Date 










































*— median chl 






grand 


mean 


chl 



























































Monthly Median Chlorophyll Anomaly, Kure 1997-2007 



0.15 



0.10 



1 0.05 

3 

r. 0.00 
(J 



-0.05 



-0.10 




<- > 

03 -D 
u -I 



CO CD CO 
^J CO CO 



03 -J 



CO CD CO 
CO CD CD 



<- > 

oi -6 

_j -i 

6 6 

o o 



03 I 

cb cb 



ro ro 



0) -g 

_3 -1 



o o o 

N. CO CO 



O 
o 

6 

co 



0) £ 
_3 -I 

o 9 



_ 6 

en en 



03 -g 
_3 -t 



o o 

01 03 03 



O 

o 

6 

(33 



> 



O O 



o 

■vl 



•median monthly chl anomaly 



Date 



Mean Seasonal Chlorophyll for Kure 1997-2007 





0.25 - 














































































0.23 - 














































































0.21 - 






















































* 












































































A 
























0.19 - 




















































/\ 






















^^ 




















































/ \ 






















_l 


0.17 - 
0.15 - 














































[_ 




I 






















3 














































. 
































M 








TV 








A 




A 








A 








A 

A 






n 




I 




A 

A\ 






A 






lf\ 








— 


0.13 - 




f\ 








' \ 








/ \ 




/ ' 








/ \ 








/ \ 




: 




\ 




\l 






/ \ 






1 










r 






_v 




r 


/ 






r 




1 










\i 






1 






1 








O 


0.11 - 


_j 






A\ 




_J I 


/A 


1 








, 




1 


\1 




j^W 




1 


I 








1 






I 


i\ 




1 






l/ y 


\ 


/ 






1 








/ 






\ 


\\ 




i " 




/ 




\ 








0.09 - 


S 
















1 






y 


\ 




















/ 






/ 


V> 




/ 






/ 




\ 














A 










,' 






£ 


v_ 




















r 






/ 


v 




/ 








/ 




\ 








n n7 








*■__<• 








^ 


• 






/ 


V 










■»- 








<_ 


/ 






/ 


\ 


*•— - 


/ 






**m* 


> 










0.05 - 
























































































































































Spr 
Fall 
Spr 
Fall 
Spr 
Fall 
Spr 
Fall 
Spr 
Fall 
Spr 
Fall 
Spr 
Fall 


Tl 

_ 


Spr 
Fall 
Spr 
Fall 
Spr 






2004 
2003 
2003 
2002 
2002 
2001 
2001 
2000 
2000 
1999 
1999 
1998 
1998 
1997 






2007 
2006 
2006 
2005 
2005 






Date 










—*—mean seasonal chl grand mean seasonal chl 








A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



0.23 



Monthly Median Chlorophyll for Special Management Area 1997-2007 






n 


c_ 
cd 


> 

T3 


c_ 

c 



n 


c_ 
cd 


> 


C_ 

c 




n 


c_ 
CD 


> 

T3 


c_ 

c 




n 


c_ 

CD 


> 

T3 


c_ 

c 




n 


c_ 
CD 


> 
■D 


c_ 
c 




n 


c_ 

CD 


> 

T3 


c_ 
c 



n 


c_ 

CD 


> 

T3 


C_ 

c 



n 


c_ 
CD 


> 

T3 


c_ 

c 




n 


c_ 

CD 


> 
T3 


c_ 
c 




n 


c_ 

CD 


> 

T3 


c_ 

c 




































-s 








- \ 
























- s 








- \ 




CO 


CO 


CD 


CD 
CO 


CD 


CO 


CO 


CD 
CD 


CO 


O 


o 


o 

o 


o 


o 


O 


O 


o 


O 


O 


O 


o 


o 


o 


O 

CO 


o 


o 


O 


O 


o 


n 


o 


o 
en 


o 


O 


o 


o 
ffi 


o 


o 


o 


o 
>l 


^l 


CO 


CO 


CO 


CD 


CD 


CO 


o 


o 


o 


-* 


- 1 




- 1 


M 


N3 


K> 


CO 


CO 


CO 


*■ 


-1^ 


-1^ 


en 


en 


en 


0) 


0) 


0) 


s 


>l 



•median chl grand mean chl 



Date 



Monthly Median Chlorophyll Anomaly, Special Management Area 1997-2007 



0.10 




o) 0.02 -I 



- 0.00 
° -0.02- 



r-t- _J ~l I rf D "7 | r-t-J-I I rf D -| | rf D -| | rf D -| | rf D "1 | r-t- D -I I rf D -i | rf D -I | 

cbcbcb¥icbcbScb66°6662°ooR66og6669666 



•median monthly chl anomaly 



Date 



Mean Seasonal Chlorophyll for Special Management Area 1 997-2007 



n 17 - 


















































































n 1^; - 


















































































"Si n 11 - 














































































2 0.11- 
W n no - 






































































































n 07 - 


































































0.05- 






n^ 


n^ 


' 


' 


' i i ' i ' i ' r» i ' i ' i ' i ' i ' i ' i ' 


, 






n 1 - 


n^ 


n^ 


r-^- 


r-^- 


i 


_i — 




i ' ■ ■ ■ ' 


r-^- 


r^ 




n 1 - 


n^ 


r-^- 


rh 



■n 


CO 


Tl 


CO 


Tl 


CO 


Tl 


CO 


Tl 


CO 


Tl 


CO 


Tl 


CO 


Tl 


CO 


Tl 


CO 


Tl 


(n 


ID 


■D 


CD 


T3 


CD 


T3 


CD 


T3 


CD 


■D 


CD 


■D 


CD 


■D 


CD 


■D 


CD 


T3 


CD 


TJ 


— 


-s 


— 


—* 


— 


- \ 


— 


- s 


— 


-5 


— 


- s 


— 


-5 


— 


- \ 


— 


- s 


— 


-5 


_i 


-I 


_i 


-i 


_i 


M 


K> 


M 


NJ 


M 


K> 


M 


M 


K> 


M 


NJ 


M 


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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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Monthly Median Chlorophyll Anomaly, Pearl and Hermes 1997-2007 





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Mean Seasonal Chlorophyll for Pearl and Hermes 1 997-2007 



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Date 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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Mean Seasonal Chlorophyll for Lisianski 1997-2007 



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■median seasonal chl grand mean seasonal chl 



Date 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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Mean Seasonal Chlorophyll for Laysan 1997-2007 





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•median seasonal chl grand mean seasonal chl 



Date 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Monthly Median Chlorophyll for Maro Reef 1997-2007 



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Date 



Mean Seasonal Chlorophyll for Maro Reef 1 997-2007 





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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

























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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Geology and Benthic Habitats 

Jonathan Weiss 12 , Joyce Miller 13 , Emily Hirsch 14 , John Rooney 1 , Lisa Wedding 56 and Alan Friedlander 67 

INTRODUCTION AND ORIGIN 

The Northwestern Hawaiian Islands 
(NWHI) are in the middle section of the 
6,126 km long Hawaiian-Emperor sea- 
mount chain, considered to be the lon- 
gest mountain chain in the world (Grigg, 
1983; Figure 3.1). Over many millions 
of years, a relatively stationary plume of 
hot mantle, or hot spot, located below 
the floor of the Pacific Plate (Grigg 1982, 
1997; Rooney et al., 2008) has and con- 
tinues to erupt at the seafloor creating 
a chain of volcanoes that comprise the 
islands, banks, atolls and seamounts 
of the Hawaiian Archipelago (Figure 
3.2). Each begins as a small subma- 
rine volcano and over time can grow to 
reach well above sea level. Eventually 
the volcanoes cool and subside as they 
slowly move away from the hot spot in a 
northwestward direction at about 8 cm/ 
yr (Clague and Dalrymple, 1987). 

The tops and edges of the volcanoes, 
if they are at or near sea level, support 
large and diverse coral reef communi- 
ties. As the volcanic edifice subsides, 
an atoll can form as reef builders keep 
the top of the volcano near sea level by 
growing vertically and creating a thick 
carbonate cap. The Darwin Point marks 
the threshold where vertical growth, or 
net accretion, of reef building organisms 
is zero or negative and the atoll drowns 
and becomes a guyot (Figure 3.3; Grigg 
et al., 2008). This point, named after 
Charles Darwin, who first proposed an 

evolutionary model for atoll formation in 1836, marks a significant milestone in the life of a Hawaiian volcano. 
Currently Kure Atoll lies very near its Darwin Point. It is the oldest Hawaiian island still above sea level although 
it consists of only 1 km 2 of emergent land and 66 km 2 of lagoon (Juvik and Juvik, 1998). Dozens of seamounts 
and guyots extend from north of Kure to the Aleutian Trench and mark the ancient remnants of volcanoes simi- 
lar to those that comprise the Hawaiian Archipelago (Davies et al., 1972). 



Figure 3.1. Oblique southern perspective of the bathymetry of the Pacific 
plate between Hawaii and the Aleutian Islands constructed to show the Ha- 
waiian-Emperor Seamount chain and the progressive subsidence of each 
volcano over time. Sources: Neall and Trewick 2008; image prepared by 
Jon Procter. 




Figure 3.2. The island of Hawaii currently sits over the hot spot that has 
formed the Hawaiian Archipelago over tens of millions of years. Photo: Ha- 
waii Volcano National Park. 



1. NOAA/NMFS/Pacific Islands Fishery Science Center, Coral Reef Ecosystem Division 

2. University of Hawaii Department of Geology and Geophysics 

3. Joint Institute for Marine and Atmospheric Research 

4. Aquatic Farms Ltd. 

5. University of Hawaii at Manoa 

6. NOAA/NOS/NCCOS/CCMA Biogeography Branch 

7. The Oceanic Institute 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



General Description of the Archipelago 
The youngest and largest of the emer- 
gent Hawaiian Islands is the island of 
Hawaii, which is composed of five vol- 
canoes including the currently active 
Kilauea Volcano and Mauna Loa and 
Mauna Kea, which are the two massive 
shield volcanoes that form the bulk of 
the island (Figure 3.4). They both extend 
from the seafloor at >5,000 m below sea 
level to >4,000 m above sea level and 
Mauna Kea is the single largest moun- 
tain on Earth. Loihi is the youngest 
submarine volcano in the archipelago 
and is located 30 km southeast of Ha- 
waii Island. Northwest of Hawaii Island 
the islands of Maui, Lanai, Kahoolawe 
and Molokai, make up the island clus- 
ter known as "Maui Nui". The highest 
point in Maui Nui is Haleakala volcano 
on Maui at 3,055 m. Further northwest 
is Oahu, the most densely populated is- 



i 




: — s 


1 




Darwin Poim 


+Q.05 j£^i nyjw 


8 


■ Subductenn 


-0.025 

Subsidence 


Uplift \ 

SubsidenteV 


E ° 

E .1 

-2 

-3 



30" 

■ North Pacific 



Figure 3.3. Reef accretion across the Hawaiian Archipelago. Note that the 
net accretion rate diminishes to zero just beyond Kure Atoll, thereby defin- 
ing a threshold for atoll formation known as the Darwin Point beyond which 
atolls drown. Source: Grigg, 1997; image: S. Hile. 



land in the chain. Kauai, the Garden Isle, is approximately 5.1 million years old and is deeply eroded with lush 
vegetation and steep cliffs (Juvik and Juvik, 1998). Niihau and Lehua, which lie 27.7 km southwest of Kauai, 
are far drier than their larger and higher elevation neighbor, Kauai. Lehua is a private island inhabited by an 
isolated population of native Hawaiians and access to the island is strictly controlled. Kaula Rock, an uninhab- 



175"W 



Pearl anf 
Hermes Atoll 

107 7\ ~ 



170°W 



165°W 



160°W 



155°W 



Midway 
Atoll 
(27.7) 



Laysan 



Gardner 
Pinnacles 



Usianski 

(23.4) 



French Frigate 
Shoals 



Nihoa Island 



Maro Reef 



'***) 



Kauai 

(7.2) Oahu 

(3.7,2.6) Molokai 
M.8> 



NDs 



Maui 
'1.3,1.1) 



Lanai, 
Kahoolawe 

(1.0) 



A 



| I Monument Boundary 



^<„ 



(0.4,0.2) 



1 75°W 



170°W 



165°W 



160°W 



155°W 



Figure 3.4. Islands, banks and atolls of the Hawaiian Archipelago. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



ited island southwest of Niihau, marks 
the transition between the main or wind- 
ward islands and the northwestern or 
leeward islands. The two thousand km 
of ocean between Kaula Rock and Kure 
Atoll separates the 10 islands and atolls 
in the NWHI and a few small emergent 
pieces of land with a total area of just 
over 15 km 2 (Juvik and Juvik, 1998). 
Northwest of Mokumanamana and Ni- 
hoa, are remnant basaltic pinnacles in- 
cluding La Perouse Pinnacle at French 
Frigate Shoals and the Gardner Pin- 
nacles. The maximum elevation of the 
islands northwest of Gardner Pinnacles 
does not exceed 12 m and on some 
atolls or islands only reaches 3 m. 

Table 3.1 shows the land areas of all 
emergent islands in the NWHI, as well 
as the submerged marine area encom- 
passed by the 10-fathom (18.29 m) and 
100-fathom (182.9 m) isobaths. Within 
10 fathoms total shallow water habitats 
constitute 1,595 km 2 , including the area 
within lagoons. The area of shallow wa- 
ter habitats within the 100 fathom isobath 
is 13,771 km 2 . However, these numbers 
are representative of a two dimensional 
surface and do not truly convey the sur- 
face area of the benthos, which, due to 
the 3-D topographic complexity of the 
seafloor, is actually substantially greater. 
Accurate bathymetric data for the NWHI 
are being collected and synthesized by 
the NOAA Coral Reef Ecosystem Divi- 
sion, the Hawaii Mapping Research 
Group, Joint Institute for Marine and At- 
mospheric Research (JIMAR), and the 
Hawaii Undersea Research Laboratory, 
at the University of Hawaii's School of 
Ocean and Earth Science and Technol- 
ogy 



Table 3.1. Area mapped by aggregated habitat cover type and geographic 
scale (km 2 ) based upon IKONOS satellite imagery. Source: NOAA, 2003. 



ISLAND 


LAND AREA (km 2 ) 


AREA 

(km 2 ) <10 

FM 


AREA (km 2 ) 
<100 FM 


Kaula 


0.6 





70.5 


Bank E of Nihoa 


Submerged 





146.9 


Nihoa 


0.7 


5.6 


570.8 


Bank SW of Nihoa 


Submerged 





336.2 


Bank NW of Nihoa 


Submerged 





63.8 


Twin Banks 


Submerged 


2.3 


72 


Mokumanamana 


0.2 


9.1 


1,557.2 


French Frigate Shoals 


0.2 


469.4 


943.4 


Southeast Brooks Bank 1 


Submerged 





29.4 


Southeast Brooks Bank 2 


Submerged 





142.3 


Southeast Brooks Bank 3 


Submerged 





158.5 


Southeast Brooks Bank 4 


Submerged 





3.4 


Brooks Bank NW of St. 
Rogatien 


Submerged 





67.8 


St Rogatien Bank 


Submerged 





383.1 


Gardner Pinnacles 


0.02 


0.7 


2,446.6 


Raita Bank 


Submerged 


16 


571.1 


Maro Reef 


Awash 


217.5 


1,935.3 


Laysan Island 


4.1 


26.4 


584.5 


Northhampton Seamounts (4) 


Submerged 





404.3 


Pioneer Bank 


Submerged 





434.6 


Lisianski Island/Neva Shoal 


1.5 


215.6 


1,246.6 


Bank NW of Lisianski 


Submerged 





106.7 


Bank SSE of Pearl and 
Hermes 


Submerged 





5.5 


Bank ESE of Pearl and 
Hermes 


Submerged 





4.8 


Pearl and Hermes Atoll 


0.3 


374.5 


816.6 


Salmon Bank 


Submerged 





163.2 


Gambia Shoal 


Submerged 





0.5 


Ladd Seamount 


Submerged 


54.2 


144.1 


Midway Atoll 


64 


85.4 


344.1 


Nero Seamount 


Submerged 


25 


71.8 


Kure Atoll 


1.0 


90.2 




Bank W of Kure 


Submerged 


NA 


NA 


TOTAL AREA 




1,591.9 


13,805.6 



Pre-Holocene Reef History 

The Hawaiian-Emperor chain includes at least 129 massive shield volcanoes that formed over the past 85 mil- 
lion years, with volcano ages generally decreasing in age towards the southeast (Jackson et al., 1975; Clague, 
1996). The overall age progression of the islands has been confirmed by several studies using radiometric 
isotopes to date volcanic rocks from islands and seamounts along the chain (Table 3.2; Clague and Dalrymple, 
1987; Garcia et al., 1987) although suitable samples for dating many of the NWHI are difficult to obtain. It has 
been proposed that the frequency of volcano formation has increased over time based on the decrease in 
volcano spacing over time. It has also been proposed that the islands at the younger end of the chain are also 
significantly higher than those formed earlier (Clague, 1996). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Table 3.2. Characteristics of the islands, atolls, and some submerged banks in the NWHI , listed in order from the north- 
west down to the southeast. Note that most of the subaerially exposed islands, sea stacks, and atolls are surrounded by 
extensive shallow banks. Island ages are from Clague (1996), with values in brackets from K-Ar dated basalt samples, 
and other ages estimated from geophysical calculations. Lagoon water volumes are from Hoeke et a/., 2006. 



ISLAND, ATOLL OR 
BANK 


TYPE OF 
FEATURE 


LONGITUDE 


LATITUDE 


AGE (MA) 


REEF 
EMER- 
GENT 
LAND 
(km 2 ) 


LAGOON 

HABITAT 

<100 m 

(km 2 ) 


BACK- 
REEF/ 
LAGOON 
VOLUME 
(10 3 m 3 ) 


SUMMIT 

DEPTH 

(m) 


Kure Atoll 


Closed atoll 


178° 19.55' 


28° 25.28' 


29.8 


0.86 


167 


141,000 


- 


Nero Seamount 


Bank 


177° 57.07' 


27° 58.88' 


29.1 


0.00 


17 


- 


68 


Midway Atoll 


Closed atoll 


177°22.01' 


28° 14.28' 


[27.7], 28.7 


1.42 


223 


213,000 


- 


Pearl and Hermes Atoll 


Closed atoll 


175° 51.09' 


27° 51.37' 


[20.6], 26.8 


0.36 


1,166 


2,930,000 


- 


Lisiankski Island Neva 
Shoal 


Open atoll 


173° 58.12' 


26° 4.2' 


23.4 


1.46 


979 


242,000 


- 


Pioneer Bank 


Bank 


173° 25.58' 


26° 0.71' 


22.8 


0.00 


390 


- 


26 


North Hampton 
Seamounts 


Bank 


172° 14.08' 


25° 26.84' 


[26.6], 21.4 


0.00 


430 


- 


5 


Laysan Island 


Carbonate 
island 


171° 44.14' 


25° 46.13' 


[19.9], 20.7 


4.11 


57 


3,600 


- 


Maro Reef 


Open atoll 


170° 38.34' 


25° 30.2' 


19.7 


0.00 


1,508 


611,000 


- 


Raita Bank 


Bank 


169° 30.04' 


25° 31.72' 


17.9 


0.00 


650 


- 


16 


Gardner Pinnacles 


Basalt sea 
stacks 


167° 59.82' 


25° 0.04' 


[12.3], 15.8 


0.02 


1,904 


- 


- 


St. Rogoties Banks 


Banks 


164° 7.26' 


24° 20.0' 


14.7 


0.00 


500 


- 


22 


Brocks Banks 


Banks 


166° 49.31' 


24° 7.03' 


[13.0], 13.6 


0.00 


320 


- 


20 


French Frigate Shoals 


Open atoll 


166° 10.75' 


23° 45.99' 


12.3 


0.23 


733 


1,910,000 


- 


Bank 66 


Bank 


165° 49.37' 


23° 51.86' 


11.9 


0.00 





- 


120 


Mokumanamana 


Basalt island 


164° 41.90' 


23° 34.64' 


[10.3], 10.6 


0.21 


1,538 


64.2 


- 


Twin Banks 


Bank 


163° 3.78' 


23° 13.08' 


8.7, 8.3 


0.00 


9.5 


- 


53 


Nihoa Island 


Basalt island 


161° 55.25' 


23° 3.73' 


[7.2], 7.3 


0.82 


246 


- 


- 



Holocene Reef Development 
Using values of island-specific cor- 
al community measures from Grigg 
(1982), vertical rates of reef accretion 
can also be estimated and are shown 
for four islands in Figure 3.5. As the fig- 
ure suggests, Grigg (1982) found that 
growth rates show a strong latitudinal 
dependence, with corals in cooler more 
northerly islands growing more slowly. 
He also noted that, based on the work of 
Gross et al. (1969), at least at the more 
northerly atolls the carbonate contribu- 
tion from coralline algae is likely to be 
significantly more than that from corals. 



:iQ 



1.65- 

1.60 - 

| 1.55 - 

1 1.50 - 

> 1.45 - 

55 

S 1.40 - 



Kure 



FFS 



Oahu 



Hawaii 



> 16 

I U 
j 12 

3 io 



.2 6 

1 4 

u 

< 2 

| 
E 



1.30 



■C 10- 

S 8- 

c 



S 6- 



5 4 " 

s 

O 2- 



l ll n li n 



Kure 



FFS 



Oahu 



Kure 



FFS 



Oahu Hawaii 




Oahu Hawaii 



Figure 3.5. Latitudinal variations in coral colony growth rates and reef ac- 
cretion across the Hawaiian Archipelago. Source: Grigg, 1982. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



BENTHIC HABITAT MAPPING 

Potential Reef Area 
Geographic information system-based 
analyses were used to derive compre- 
hensive, consistent estimates of the 
potential area of broadly defined, shal- 
low-water, tropical and subtropical coral 
ecosystems within the territorial sea and 
exclusive economic zone of the United 
States (Rohmann et al., 2005). Nautical 
charts, published by NOAA's Office of 
the Coast Survey, provide a consistent 
source of 10-fathom (ca.18 m) and 100- 
fathom (ca. 183 m) depth curve infor- 
mation. The 10-fathom or 100-fathom 
depth curves are used as surrogates for 
the potential distribution and extent of 
shallow-water coral ecosystems in trop- 
ical and subtropical U.S. waters (Fig- 
ure 3.6). The NWHI constitute between 
4.3% and 12.6% of the total U.S. poten- 
tial coral reef ecosystem area within 10 
fathoms, depending on inclusion versus 
exclusion of the west Florida shelf area 
(Rohmann et al., 2005). The NWHI con- 
stitute 9.6% of the total U.S. potential 
coral reef ecosystem area within 100 
fathoms, with the inclusion of the west 
Florida shelf area (Rohmann et al., 
2005). 

NOAA Mapping Programs 

In support of the U.S. Coral Reef Task 
Force's mission to "Produce compre- 
hensive digital maps of all shallow (<30 
m) coral reef ecosystems in the United 
States and characterize priority mod- 
erate-depth reef systems by 2009," 
NOAA has developed a comprehensive 
mapping program in the Pacific Region 
using IKONOS satellite imagery in 
shallow water (<30 m) and multibeam 
sonar technology in depths as deep 
as 3,000 m (Table 3.3). In intermediate 
depths (10-30 m) IKONOS and multi- 
beam mapping techniques can provide 
complementary or overlapping cover- 
age. In addition, IKONOS images can 
be used to create "estimated depths" 
to fill bathymetric gaps in very shallow 
water (<15 m) where multibeam ves- 
sels cannot safely survey (Stumpf and 
Holderied, 2003). 




Figure 3.6. Potential reef area from the shoreline to 10-fathoms (top) and 
100-fathoms (bottom) based on NOAA nautical charts. Source: Rohmann 
etal.,2005. 

Table 3.3. Area in the NWHI mapped by aggregated habitat cover type 
based upon IKONOS satellite imagery. Source: NOAA 2003. 



AGGREGATED HABITAT COVER TYPE 


AREA 
MAPPED 


PERCENT 
TOTAL 


Hardbottom with >10% live coral 

Hardbottom with >10% crustose coralline algae 


108.8 


4.61 


7.3 


0.31 


Hardbottom (uncolonized) 


101.4 


4.30 


Hardbottom with >10% macroalgae 
Hardbottom with indeterminate cover 


105.2 


4.46 


822.8 


34.85 


Unconsolidated with 10% or less macroalgae or 
seagrass 


1,149.6 


48.70 


Unconsolidated with >10% macroalgae or 
seagrass 

Total Habitat Area Classified 


65.8 


2.79 


2,360.8 


100.00 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Shallow Water IKONOS Satellite Mapping 
NOAA's Biogeography Branch has 
sponsored a shallow water (0-30 m) 
benthic habitat mapping program us- 
ing IKONOS satellite imagery, which in 
2003 produced the Atlas of the Shallow- 
Water Benthic Habitats of the North- 
western Hawaiian Islands (available at 
http://ccma.nos.noaa.gov/ecosystems/ 
coralreef/nwhi/welcome.html). IKONOS 
high-resolution satellite imagery was 
used to derive benthic habitat maps, 
estimated depth, and the color images. 
The detailed benthic habitat classifica- 
tion scheme was designed to categorize 
benthic habitat by substrate category 
(e.g., unconsolidated and hardbottom), 
structure (e.g., linear reef or pavement) 
and cover (e.g., coral or macroalgae; 
Figure 3.7, Table 3.4; http://biogeo.nos. 
noaa.gov). Of the area mapped, 49% 
was unconsolidated sediment while 
35% was indeterminate. 



3% 



5% 



49% 




35% 



■ Hardbottom with >10% live 
coral 

■ Hardbottom with >10% 
crustose coralline algae 

3 Hardbottom (uncolonized) 



■ Hardbottom with >10% 
macroalgae 

■ Hardbottom with indeterminat 
cover 

D Unconsolidated with 10% or 
less macroalgae or seagrass 

D Unconsolidated with >10% or 
less macroalgae or seagrass 



Figure 3. 7. Area in the NWHI mapped by aggregated habitat cover type 
based upon IKONOS satellite imagery. Source: NOAA, 2003. 



Table 3.4. Area mapped by aggregated habitat cover type and geographic scale (km 2 ) based upon IKONOS satellite 
imagery. Source: NOAA, 2003. 



TOTAL 


KURE 
ATOLL 


MIDWAY 
ATOLL 


PEARL & 

HERMES 

ATOLL 


LISIANSKI 
ISLAND 


LAYSAN 
ISLAND 


MARO 
REEF 


FRENCH 
FRIGATE 
SHOALS 


MOKUMANA- 
MANA 


NIHOA 
ISLAND 


Hardbottom with >10% 
live coral 


1.8 


1.4 


20.3 


16.4 


5.8 


14.8 


48.3 





0.1 


Hardbottom with >10% 
crustose coralline algae 


0.7 


0.1 








0.5 


1.3 


4.7 








Hardbottom 
(uncolonized) 


11.6 


14.9 


13.7 


0.9 


2.9 


6.8 


49.9 





0.7 


Hardbottom with >10% 
macroalgae 


5.8 


22.4 


62.2 


6.1 


0.1 


0.4 


3.7 





4.5 


Hardbottom with indeter- 
minate cover 


8.4 


6.7 


49.3 


183.5 


81.7 


180.1 


46.1 


208.1 


58.9 


Unconsolidated with 
10% or less macroalgae 
or seagrass 


38.8 


49.9 


226.2 


231.8 


36.2 


295.7 


241.5 


19.5 


10 


Unconsolidated with 
>10% macroalgae or 
seagrass 


2.7 


0.2 


19.9 








19.6 


23.4 








Total Habitat Area 
Classified 


69.8 


95.5 


391.6 


438.7 


127.2 


518.7 


417.6 


227.6 


74.1 



Moderate-Depth Multibeam Mapping 

NOAA's Coral Reef Ecosystem Division (CRED) initiated a moderate-depth multibeam mapping program 
which was conceived in 2001, implemented between 2002 and 2005, and has produced over 45,000 km 2 of 
bathymetric data in the NWHI since 2002 (Table 3.5; Miller et al., 2003). This mapping program is designed to 
extend and be complementary to the shallow-water IKONOS mapping program discussed above. 



In 2002 multibeam surveys to define 25, 50, and 100-fm isobaths in the NWHI were conducted by NOAA 
and University of Hawaii (UH) personnel aboard UH's R/V Kilo Moana, using Kongsberg/Simrad EM1002 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 3.5. NWHI multibeam mapping statistics and estimates. 





MAPPING COMPLETED 
2002-2008 


ESTIMATE TO 
COMPLETE 


km 2 


Days 


Days Remaining 


Deep (100-5,000 m) 


41,664 


28 


67 


Mid-Depth (10-100 m) 


3,854 


134 


275 


Totals 


45,518 


162 


342 



the banks and atolls that are shown in 
Figure 3.8. Moderate depth multibeam 
sonar surveys were conducted in the 
NWHI between 2003 through 2008 by 
personnel from CRED, NOAA's Office 
of National Marine Sanctuaries, and 
other partners using mapping systems 
aboard the NOAA Ship Hiialakai and 
the survey launch R/V Acoustic Habitat Investigator (AHI). The Hiialakai is equipped with two Kongsberg/ 
Simrad multibeam sonars: a 30-kHz EM300 with mapping capability from approximately 100-3000+ m and a 
300-kHz EM3002D with mapping capability from about 5-150 m. The R/V AHI has a 240-kHz Reson 8101ER 
with mapping capability from about 5-300 m. Both vessels have Applanix POS/MV motion sensors, which pro- 
vide navigation and highly accurate readings of the vessel motion in all axes. Optical validation data have also 
been collected since 2001 using towed and drop camera systems aboard the Hiialakai, AHI and the NOAA 
Ship Oscar Elton Sette. 

Bathymetric data from these 2003-2008 Hiialakai and AHI surveys add to previously published data (Miller et 
al., 2003) from the 2002 R/V Kilo Moana surveys as well as estimated depths from IKONOS imagery Figure 
3.8 and Table 3.5 show current bathymetric coverage in NWHI. 



Lisianski to Maro 



\ * 



@- 



Bathymetry <6D mj 

Multibeam & 
IKONOS-derlved 



■I KK50 A 

15 30 » 

^^K=^^^^MP Ida ronton 



<5 



Q> 



HffW «?W 



Nihoaand 
southward 




!': Il -.■•■'. -:t .- i'.H in 

Multibeam & 
IKONOS -derived 

Dapllt i rn : 

N 
HI tO S3 





^*. 






IfJ-W U2-W 



171-w WW 





Kure to Lisianski 




^ 9 , 




"&>+**■ 


S Si'.' V ■">■■.' V ■■■"'• II.' 

Multlbtam $ 
iKONOS^erlwed 






Depth (m> 






1 HJ A0 B0 

^h=^^^h K>k.iiv.|vii 







it*-w ire-w mrvt i7i*w rjvvt mm 



Gardner to 
Mokumanamana 




Figure 3.8. Multibeam maps of all data collected in the NWHI through 2006. Maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Bathymetric grids at various resolutions are updated annually and published on the web at the Pacific Islands 
Benthic Habitat Mapping Center (PIBHMC; www.soest.hawaii.edu/pibhmc). As shown in Figure 3.8, some ba- 
thymetric data have been collected and processed at all of the islands and banks in the NWHI in water depths 
ranging from 3 to 3,000 m with almost complete coverage at Kure, Midway, Pearl and Hermes, Brooks Banks 
and French Frigate Shoals, and partial coverage at other locations. The bank on the southwest side of French 
Frigate Shoals was the first area to be mapped in early 2005 and 95% completed in 2008, and this data set is 
used later in this chapter to illustrate the various benthic habitat mapping products, their potential uses, and 
interpretation. Similar products for other banks are regularly added to the PIBHMC web site as mapping, data 
processing, product and metadata generation, and interpretation are completed. 

The geomorphological data layers of substrate, slope, rugosity, and bathymetric position index (BPI) produced 
at the PIBHMC are derived from multibeam bathymetry. Derivative data products (e.g., slope, rugosity and 
BPI) add geomorphological information about characteristics (e.g., roughness) that may assist in determining 
benthic habitat utilization. An explanation of each derivative type is given here. At this time a complete set of 
derivate products has been developed only for French Frigate Shoals. 

Rugosity: Cell values reflect the surface area to planimeteric area ratio (surface area) / (planimetric area) 
for the area contained within that cell's boundaries. This measure provides an index of topographic rough- 
ness and convolutedness (Jenness, 2003). Distributions offish and other mobile organisms are often found 
to positively correlate with increased complexity of the seafloor. Investigations are underway for the de- 
velopment of the most appropriate spatial metrics for quantifying benthic complexity for the purposes of 
relating these metrics to fish distributions in Pacific coral reef ecosystems. Results of the Jenness (2003) 
method are provided as a standardized and well-documented interim product. 

Slope: Cell values reflect the maximum rate of change (in degrees) in elevation between neighboring 
cells. 

Substrate: This is a preliminary product that is still under development. Cell values reflect whether the 
seafloor is hard bottom or soft bottom based on an unsupervised classification run in Environment for Vi- 
sualizing Images (ENVI) software. The classifications (hard bottom versus soft bottom) are based on back- 
scatter, bathymetry, acoustic derivatives and optical data. 

Bathymetric Position Index: BPI is a second order derivative of bathymetry. The derivation evaluates 
elevation differences between a focal point and the mean elevation of the surrounding cells within a user 
defined annulus or circle. A negative value represents a cell that is lower than its neighboring cells (depres- 
sions) and a positive value represents a cell that is higher than its neighboring cells (crests). Larger num- 
bers represent more prominent features on the seafloor, which differ greatly from surrounding areas. Flat 
areas or areas with a constant slope produce near-zero values. (Lundblad et al., 2006). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Summary of Multibeam Data 
Based on available multibeam data, summary 
statistics (slope, aspect ratio and rugosity) were 
developed for five depth bins based on natu- 
ral breaks (9-226 m, 226-588 m, 588-1,045 m, 
1,045-1,649 m and 1,649-3,071 m). Slope values 
overall are lowest in the shallow (<226 m) depth 
bin (Figure 3.9). Islands further upchain such as 
Kure, Midway, Pearl and Hermes and Lisianski 
have higher slope values, on average, compared 
with locations further southeast along the chain. 
Aspect ratio (A Z/AX) does not vary greatly by 
depth bin or among reefs. Rugosity increases 
slightly with depth and the highest rugosity was 
found at Pearl and Hermes Atoll. 



Island Profiles 

The following sections summarize the geologic 
and benthic habitat information available for each 
emergent island in the NWHI, as of January 2009. 
The data include results from both IKONOS satel- 
lite imagery and multibeam data analyses. 




■ NIH 
DMMM 

■ FFS 

□ MAR 

□ LIS 

□ PHR 

■ MID 

□ KUR 



■ NIH 

□ MMM 

■ FFS 

□ MAR 

□ LIS 

□ PHR 

■ MID 

□ KUR 



■ NIH 

□ MMM 

■ FFS 

□ MAR 

□ LIS 

□ PHR 

■ MID 

□ KUR 



Figures 3. 9. A) Average slope by depth bin from multibeam so- 
nar data collected at select islands within the NWHI; B) Average 
aspect ratio by depth bin from multibeam sonar data collected at 
select islands within the NWHI; and C) Average rugosity by depth 
bin from multibeam sonar data collected at select islands within 
the NWHI. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Nihoa Island 

Nihoa Island is located approximately 
249.4 km northwest of Kauai, the clos- 
est to the Main Hawaiian Islands (MHI). 
Measuring roughly 0.68 km 2 , this island 
is the largest emergent volcanic island 
within the Monument and the tallest, 
reaching an elevation of 275.2 m at 
Miller Peak. It is also the geologically 
youngest island within the Monument, 
with an age calculated at 7.3 million 
years (Clague, 1996). Nihoa is a deeply 
eroded remnant of a once large volcano, 
and the large basaltic shelf of which it is 
a part stretches 28.9 km in a northeast- 
southwest direction and ranges between 
34.1 and 66.1 m deep (NOAA, 2003). 
The island's two prominent peaks and 
steep sea cliffs are clearly visible from a 
distance, rising like a fortress above the 
sea. The island's northern face is com- 



i% 



13%- 



6% 




80% 



I Hardbottom with >10% live coral 



■ Hardbottom with >10% crustose 
coralline algae 

□ Hardbottom (uncolonized) 



□ Hardbottom with >10% 
macroalgae 

□ Hardbottom with indeterminate 
cover 

□ Unconsolidated with 10% or less 
macroalgae or seagrass 

■ Unconsolidated with >10% 
macroalgae or seagrass 



Figure 3.10. Benthic habitats around Nihoa 
data. Source: NOAA, 2003. 



based on IKONOS satellite 







i 

N 

i O 0.3 06 

F\ m Klomalan 


^^ 




B • 1 1 1 c HsbttAt ; v i i - 



~-jH«a»Hm 



; Uncpnt43i|« 



101 i4-Jfi"W 




wwww 




Figure 3.11. IKONOS satellite image (top left), benthic habitat map (top right), extent of water depth <20 m (bottom left), 
and habitat zones for Nihoa Island (bottom right). Maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

posed of a sheer cliff made up of successive layers of basaltic lava, within which numerous volcanic dikes are 
visible. The island's surrounding submerged reef habitat totals approximately 574.6 km 2 and is a combination 
of uncolonized hard bottom, macroalgae, pavement with sand channels and live coral, and uncolonized volca- 
nic rock (NOAA, 2003; Figures 3.10, 3.11). The principal shallow water bottom habitats around Nihoa consist 
of hard basalt as vertical walls, horizontal wave-cut basalt benches, elevated mounds, and large blocks and 
boulders. Nihoa supports coral communities with very limited total habitat, most of which is not protected from 
the heavy and chronic wave action that strikes this small island from all directions. These habitats have been 
shaped by and are constantly eroded by the pounding waves. 

Multibeam surveys were conducted around Nihoa Island and on West Nihoa Bank on several different cruises, 
including R/V Kilo Moana KM-02-06, Hiialakai HI-05-01 and Hiialakai HI-06-12. The first two survey patterns 
around Nihoa Island were designed to delineate the 25-, 50-, and/or 100-fm boundaries needed for the Na- 
tional Marine Sanctuary designation process. Slope, aspect, and rugosity all increased with depth although the 
difference was small (Table 3.6). In Tables 3.6 -3.13 minimum, maximum, range, mean and standard deviation 
for bathymetry are in meters; slope and aspect are in degrees (0-360); and rugosity is a dimensionless ratio 
of surface area to planimetric area. High rugosity typically indicates a rough and complex substrate that often 
correlates with potential coral habitat. 

Bathymetry data for all islands was merged into a single raster. This was reclassified into five depth classes 
using natural breaks - a method which seeks to equalize the variation between each class - in the ArcGIS 9.2 
Spatial Analyst extension. Using these 5 classes as zones, zonal statistics were run on bathymetry, slope, 
aspect and rugosity by island, using the "zonal statistics as table" tool in Spatial Analyst. This resulted in 



Table 3.6. Summary statistics for multibeam surveys conducted around Nihoa Island and on 


West Nihoa Bank. 


NIHOA 


DEPTH CLASS 


AREA 


MINIMUM* 


MAXIMUM* 


RANGE 


MEAN 


STANDARD 
DEVIATION 


Bathymetry 


9 to 226 


353.68 


-225.17 


-25.21 


199.96 


-69.46 


41.34 


266 to 588 


248.65 


-500.00 


-225.17 


274.82 


-356.89 


65.18 


Slope 


9 to 226 


- 


0.00 


68.01 


68.01 


3.72 


7.79 


266 to 588 


- 


0.00 


75.99 


75.99 


8.96 


7.66 


Aspect 


9 to 226 


- 


0.00 


360.00 


360.00 


180.02 


111.23 


266 to 588 


- 


-1.00 


360.00 


361.00 


191.13 


118.97 


Rugosity 


9 to 226 


- 


1.00 


2.86 


1.86 


1.01 


0.06 


266 to 588 


- 


1.00 


4.44 


3.44 


1.03 


0.05 


*NOTE: the minimum represents the minimum value of a given metric within a universal depth class; maximum represents the maxi- 
mum value of a given metric within a universal depth class. 



Bathymetry (5 mh 
Mullibsam data 


DejJlhim) 




■1 It 




■H -'' 


N 


2 * a r% 






Bathymetry (29 m) 
Mullibaam data 



L " d A 

4 a '^ 



Figure 3.12. 5 m and 20 m bathymetry for Nihoa with derived depths from IKONOS imagery near island center. Maps: L. 
Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



f 





C 



J 



Slope {5 mh 
Multibsam data 
Slope 
^H vary sum> ut 

i 



' MbfrflfltO" 
1 bud 



Figure 3.12 (continued). Rugosity (5 m), and slope (5 m) for Nihoa with derived depths from IKONOS imagery near island 
center. Maps: L. Wedding. 



the generation of summary statistics for 
each island in terms of the NWHI as a 
whole, making it possible, for example, 
to compare the average slope of Kure 
with that of Maro within the same depth 
class. 

Cruise Hiialakai HI-06-12 surveys on 
West Nihoa Bank were conducted to 
provide continuous coverage in order 
to better delineate topography on sub- 
merged banks; the resulting maps re- 
vealed some intriguing features on the 
southern part of West Nihoa Bank (Fig- 
ure 3.12). Surveys of bottom fish habi- 
tats (Figure 3.13) and fish abundance 
were also conducted during Hiialakai 
HI-06-12. 

Mokumanamana Island 
Mokumanamana Island is a hook- 
shaped dry volcanic island that includes 
about 18 hectares of land with 9.1 km 2 
of potential coral reef habitat within 10 
fathoms and a large bank that includes 
1,557 km 2 of habitat within 100 fathoms. 
Mokumanamana is a dry volcanic island 
shaped like a fishhook, and includes ap- 
proximately 0.18 km 2 of land. Geologists 
believe the island, with an estimated age 
of 10.6 million years, was once the size 
of Oahu in the MHI, with a maximum 
paleo-elevation of 1,036 m (Clague, 
1996), but due to centuries of erosion 
its highest point, at Summit Hill, is now 
only 84.1 m above sea level. All shallow 



162"t2 , «'W 








I I I f I T~~ T 



Nihoa hi. : i ! .i Vj- T;-"r ! -y 




t&r-iz^-w 



Figure 3.13. Detailed hillshade of bathymetric data on the southern portion 
of West Nihoa bank. 




IHardbottom with >10% live coral 



■ Hardbottom with >10% crustose 
coralline algae 

□ Hardbottom (uncolonized) 



□ Hardbottom with >10% macroalgae 



□ Hardbottom with indeterminate 
cover 

□ Unconsolidated with 10% or less 
macroalgae or seagrass 

■ Unconsolidated with >10% 
macroalgae or seagrass 



91% 



Figure 3.14. Percent composition of mapped benthic habitats at Mokuma- 
namana based on NOAA benthic habitat maps. Source: NOAA, 2003. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



rapcrarw 




iwwa r w 



1W4t , I4 , W 



164 t 42 , JB"W 





T^kc 


• ' ■ . -'■- 




N 

Land y 
_ _ -_„ Q 0.1 0.2 D.4 fi 













benthlc Kn bitat Type 



CD 



urn 

0-B 1 



A 



lU'tt'afA 



191 HWVf 




164 , 43 , irW 



■Bfwwrw 



Figure 3.15. IKONOS satellite image (top left), benthic habitat map (top right), extent of water depth <20 m (bottom left), 
and habitat zones (bottom right) for Mokumanamana. Maps: L. Wedding. 

Table 3. 7. Summary statistics for multibeam surveys conducted around Mokumanamana in 2002. 



MOKUMANAMANA 


DEPTH CLASS 


AREA 


MINIMUM* 


MAXIMUM* 


RANGE 


MEAN 


STANDARD 
DEVIATION 


Bathymetry 


9 to 226 


202.99 


-286.14 


-1.03 


285.11 


-90.93 


57.43 


266 to 588 


362.93 


-737.97 


-145.73 


592.23 


-376.12 


88.38 


588 to 1,046 


77.04 


-1,132.92 


-394.30 


738.62 


-812.10 


125.46 


1,046 to 1,649 


53.09 


-1,500.00 


-953.00 


547.00 


-1,290.05 


140.25 


Slope 


9 to 226 


- 


0.00 


69.72 


69.72 


7.81 


10.13 


266 to 588 


- 


0.00 


77.49 


77.49 


8.38 


8.40 


588 to 1,046 


- 


0.00 


77.45 


77.45 


13.91 


10.48 


1,046 to 1,649 


- 


0.00 


76.94 


76.94 


20.04 


13.80 


Aspect 


9 to 226 


- 


-1.00 


360.00 


361.00 


179.46 


113.87 


266 to 588 


- 


-1.00 


360.00 


361.00 


168.49 


120.76 


588 to 1,046 


- 


-1.00 


360.00 


361.00 


149.07 


119.73 


1,046 to 1,649 


- 


-1.00 


360.00 


361.00 


167.43 


126.94 


Rugosity 


9 to 226 


- 


1.00 


3.33 


2.33 


1.03 


0.09 


266 to 588 


- 


1.00 


5.67 


4.67 


1.03 


0.07 


588 to 1,046 


- 


1.00 


4.51 


3.51 


1.07 


0.13 


1,046 to 1,649 


- 


1.00 


4.87 


3.87 


1.16 


0.25 


*NOTE: the minimum represents the minimum value of a given metric within a universal depth class; maximum represents the maxi- 
mum value of a given metric within a universal depth class. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

marine habitats are basalt surfaces exposed to high wave action and the effects of scour (surge combined with 
sand and other sediments) is evident from the wave-cut bench in West Cove and the deeply cut sand channels 
and chasms at several locations in deeper water (Figures 3.14, 3.15). Reef growth in shallow waters, if any, is 
minimal and the punishing effects of large waves as demonstrated by the high wave cut sea cliffs above sea 
level and wave planed benches and shelves below sea level. The bank provides excellent habitat for spiny 
lobsters (Panulirus marginatus) and slipper lobsters (Scyllarides squammosus), especially in habitats of less 
than 27.4 m depth and high benthic relief (Parrish and Polovina, 1994). 



Multibeam surveys around Mokumana- 
manawere conducted in 2002 (KM0206) 
and 2008 (HI0804; Table 3.7, Figure 
3.16). The 2002 surveys were for 25-, 
50-, and/or 100-fm boundary delineation 
and the survey in 2008 was planned to 
better delineate the Necker Ridge that 
runs southwest from Mokumanamana, 
which is under consideration as a pos- 
sible extension to the U.S. Exclusive 
Economic Zone. The data from the 
2008 surveys have not yet been fully 
processed and Figure 3.16 shows only 
data collected in 2002. Slope and rugos- 
ity increase with increasing depth while 
the aspect ratio is highest in the shallow- 
est depth range (<226 m; Table 3.7). 




Figure 3.16. Multibeam bathymetric data around Mokumanamana Island 
collected in 2002 and 2006 derived depths from IKONOS imagery near 
island center. Map: L. Wedding. 



6% 



12% 



12% 



IHardbottom with >10% live coral 



■ Hardbottom with >10% crustose 
coralline algae 

□ Hardbottom (uncolonized) 



French Frigate Shoals 
French Frigate Shoals is the largest atoll 
in the chain, taking the form of an 28.9 
km long crescent. It is estimated to be 
12.3 million years old (Clague, 1996). 
The shoals consist of 0.27 km 2 of total 
emergent land surrounded by approxi- 
mately 931 km 2 of coral reef habitat, with 
a combination of sand, rubble, uncolo- 
nized hard bottom, and crustose coral- 
line algae in the windward and exposed 
lagoon areas, and patch and linear coral 
reefs in more sheltered areas (NOAA, 
2003; Figures 3.17, 3.18). Tern Island in 
the atoll is the site of a U.S. Fish and 
Wildlife Service field station, which occu- 
pies a former U.S. Coast Guard (USCG) 
Long-Range Aids to Navigation (LO- 
RAN) station that closed in 1979. The 
lagoon is also unusual in that it contains 
one exposed volcanic pinnacle (La Per- 
ouse) representing the last vestiges of 
the high island from which the atoll was 

derived, as well as approximately nine low, sandy islets. The sand islets are small, shift position, and disappear 
and reappear. 




n% 



57% 



1% □ Hardbottom with >10% 
macroalgae 

□ Hardbottom with indeterminate 
cover 

□ Unconsolidated with 10% or less 
macroalgae or seagrass 

■ Unconsolidated with >10% 
macroalgae or seagrass 



Figure 3.17. Percent composition of mapped benthic habitats at French 
Frigate Shoals based on NOAA benthic habitat maps. Source: NOAA 
2003. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 





ferlh r /inhmi lypii 
^| ri:r.JLt~l!<-*-, ■ vj . tnnk 



a 25 5 



A 





*^ 















iwzj'w 166'ifl'w 




Figure 3.18. IKONOS satellite image (top left), benthic habitat map (top right), extent of water depth <20 m (bottom left), 
and habitat zones (bottom right) for French Frigate Shoals. Maps: L. Wedding. 



Table 3.8. Summary statistics for multibeam surveys conducted around French Frigate Shoals (2002-2008). 



FRENCH FRIGATE SHOALS 



Bathymetry 



Slope 



Aspect 



Rugosity 



DEPTH CLASS 

9 to 226 



266 to 588 



588 to 1,046 
9 to 226 



266 to 588 



588 to 1,046 
9 to 226 



266 to 588 



588 to 1,046 
9 to 226 



266 to 588 



588 to 1,046 



AREA 

681.15 



284.10 



54.56 



MINIMUM* 

-294.50 



MAXI- 
MUM* 

0.00 



-620.51 



-134.19 



-699.90 
0.00 



-572.38 
70.49 



0.00 



73.04 



0.00 
-1.00 



62.14 
360.00 



-1.00 



360.00 



-1.00 
1.00 



359.99 
3.25 



1.00 



3.74 



1.00 



2.29 



RANGE 

294.50 



486.32 



127.52 
70.49 



73.04 



62.14 
361.00 



361.00 



360.99 
2.25 



2.74 



1.29 



MEAN 

-42.15 



-400.26 



-638.08 
2.40 



6.28 



6.84 
150.34 



171.14 



235.36 
1.01 



1.01 



1.02 



STANDARD 
DEVIATION 

50.69 



100.79 



33.01 
4.33 



6.10 



5.45 
115.05 



127.29 



79.11 
0.03 



0.03 



0.04 



*NOTE: the minimum represents the minimum value of a given metric within a universal depth class; maximum represents the maxi- 
mum value of a given metric within a universal depth class. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 





/^ 


V' <v , , v ^ 




I 




BtWiymetry 
MulUbaain data (S m> 

Dapffi in: 

N 

■ •™ A 

1 2 i 


v 







5"3im lES'SCW 



Slope [5 H) 

Mullibsam daU 



I Ucr^SiccnJo' 
' *iyFtoH>" 



■nlw^airn A 




Figure 3.19. 5 m (top left) and 20 m bathymetry (top right), rugosity (5 m)(bottom left), and slope (5 m)(bottom right) for 
French Frigate Shoals. 20-m bathymetry plot includes derived depths from IKONOS imagery. Maps: L. Wedding. 

Slope increases rapidly between the shallow and intermediate depth range (266-588 m) and then increases 
slightly between 588 and 1,046 m. Aspect ratio shows a gradual increase from shallow to deep depth bins 
while rugosity did not vary with depth (Table 3.8). 

French Frigate Shoals is the first island for which a substrate type map has been produced. Using depth, back- 
scatter, multibeam derivatives such as rugosity, slope, and variance (Figure 3.19), an unsupervised classifica- 
tion was performed to classify the substrate type into hard and soft bottom classes (Figure 3.20). Figures 3.21 
through 3.24 shows information used in developing hard/soft and BPI maps. These products are designed to 
aid management agencies in developing sampling protocols that focus on hard (non-sand) substrates for coral 
benthic habitat studies and benthic habitat maps. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



n: — 



166'21'0"W 
,. .-I 



1B6°18'0"W 

T7^ 



lee-is'O'w 

i „** I? ■ - - * 



1B6°12 , 0"W 




n_ru i km 

05 1 2 



n_ru 1 km 

0.5 1 2 



T-TT 



BATHYMETRY (m) 
■ ' I I I 



BACKSCATTER MAGNITUDE 



: 



-50-100 -2M 



-i.;li 



Figure 3.20. (A) Multibeam bathymetry data collected on the bank top at French Frigate Shoals. Black box corresponds 
to the location of the bathymetry (B) and backscatter (C) close-ups. Stars in (B) and (C) indicate the locations, from top to 
bottom (northwest to southeast), of the TOAD frame grabs shown in (D, E, and F). These data are used to create accurate 
habitat maps for moderate depth ecosystems in the NWHI using image processing techniques. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



166°24 P W 



Substrate Type (S m ) 
Unsupervised class. 

| Hard Bottom 
^\ Soft Bottom 

Land 

Water <20 m 

12 4 

■ Kilometers 



166"24'W 



166°1S'W 



166°12'W 



166 a 6-W 




166"18'W 



166°12'W 



166"6'W 



Figure 3.21. Hard/soft substrate type from French Frigate Shoals as produced by an unsupervised classification using 
ENVI. 



166'24'W 



166°18W 



ibs'^'w 



BPi Zones 
Multibeam data (5 m) 

| C rests 

| Depressions 

^J Flals 

| Slopes 

^Land 

Water <20 m 
12 4 



N 

A 



■ Kilometers 




166°24'W 



166°18'W 



166°12'W 



Figure 3.22. Bathymetric Position Index (BPI) zones for French Frigate Shoals. Source: Lundblad et a/., 2006. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



BPI Structures 


Multibeam data (S m) 


□ 


Narrow Depression 


^| 


Depression on Ftat 


CCI 


Midslope Depression 


CCI 


Depression on Crest 


§■ 


Open Depression 


CC 


Broad Flat 


CC 


Shelf 


■1 


Escarpmeni 


CD 


Crest in Depression 


^H 


Crest on Flat 


CC 


Midslope Cresl 


^| 


Narrow Crest 


■1 


NearVertical Wall 


1 


Land 

N 
Water <20 m * 

2 4 ^ 

zz^^^h Kilometers 




Figure 3.23. Bathymetric Position Index (BPI) structures for French Frigate Shoals. Source: Lundblad et al., 2006. 




Figure 3.24. French Frigate Shoals backscatter (top) and optical data (center) used to develop hard soft maps. The bot- 
tom panel shows a profile of the terrain from concurrent bathymetric data. Red dots indicate location of three photographs. 
Dark areas in the backscatter indicate high intensity and can often be correlated to areas of coral cover, hard substrate, 
and elevated bathymetry (two left photos), while lighter areas can often be correlated with sandy, softer substrate and 
depressions. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Brooks and St. Rogatien Banks 

The Brooks and St. Rogatien Banks include four submerged banks between French Frigate Shoals and Gard- 
ner Pinnacles (Figure 3.25). These four banks are guyots (flat-topped seamounts) and all have at least two 
different terraces near their summits that indicate previous sea level stands. 




B 



1BB J5MV 



Brooks Bank.NWJH I 

WWW Ski Mi..it-k ■■ 
MJtftRHm EallijiraUy 
lOmriiraiTni 




aMmwtTfcrin* 




ID Peispecuvy view of Brooks Bank, NWHI looking From MS oeafees. Vt - Bt 



Figure 3.25. 3-D perspective of Brooks Banks. (A) Perspective view of the Brooks Banks looking from the northeast. The 
Brooks Banks exhibit a classic flat-topped morphology created by erosion when the banks were previously at or near sea- 
level. Multiple terraces around the bank edges are evidence for additional sea-level stands (B,C). Submarine canyons 
incise the steep bank edges and blocks of material at the base of the slopes are probably slumps or landslide deposits. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Gardner Pinnacles 

Gardner Pinnacles consists of two emergent basaltic volcanic peaks estimated to be 15.8 million years in age 
(Clague 1996), which represent the oldest high islands in the Hawaiian chain. In scale, these pinnacles are 
small, the largest reaching only 54.8 m high and having a diameter of approximately 179.8 m. Due to their 
limited size, they support only a single species of land plant (Portulaca lutea) and a few terrestrial arthropod 
species, but they are by contrast excellent habitat for seabirds (Clapp, 1972). Guano from such seabirds gives 
the peaks a "frosted" appearance, indicating their importance as roosting and breeding sites for at least 12 
subtropical species. These remnant volcanic pinnacles are surrounded by approximately 2,428 km 2 of coral 
reef habitat, most of which is in waters 18.3 m or deeper (Figure 3.26). The shallow water reef area within 10 
fathoms covers less than one square kilometer (0.7 km 2 ) but the surrounding bank out to 100 fathoms cov- 
ers 2,428 km 2 . The relatively flat bank is in the 30 to 40 m depth range and consists of mostly sand and algal 
bottom with occasional rock outcroppings. The Pinnacles do not offer much protection from heavy waves and 
corals are more abundant on elevated surfaces and behind rises or mounds that are protected from wave ac- 
tion. The lack of shallow water environments limits the number of reef building species that can survive the 
conditions at the reefs and powerful wave action reduces the growth rate of corals (Grigg 1981), coralline algae 
and other reef-building organisms. 

Multibeam data were collected in 2002 at Gardner Pinnacles on cruise KM0206 on the 25, 50 and/or 100-fm 
isobaths in order to delineate boundaries for the National Marine Sanctuaries program (Figure 3.27). 






,A 



Figure 3.26. IKONOS satellite image (top left), extent of water depth <20 m (top right), and habitat zones (bottom left) for 
Gardner Pinnacles. Maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



168°40'W 



168°20'W 



168°O r W 



■\67°W\N 



Bathymetry 
Multibeam data (20 m] 

Depth (m) 



-1000 



N 

A 



4 8 16 

Kilometers 




o 

CM 



_o 



168°4ctw 



168°20 , W 



les^cw 



167°40 , w 






Figure 3.27. Gardner Pinnacles multibeam data collected in 2002 during KM0206. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Maro Reef 

Maro Reef is a largely submerged open 
atoll 19.7 million years old (Clague, 
1996). At very low tide, only a small 
coral rubble outcrop of a former island 
is believed to break above the surface; 
as a result, Maro supports no terrestrial 
biota. In contrast, the shallow water reef 
system is extensive, covering nearly 
2,023 km 2 , and is the largest coral reef 
in the Monument. Maro's reefs are intri- 
cate and reticulated, forming a complex 
network of reef crests, patch reefs, and 
lagoons. Deepwater channels with irreg- 
ular bottoms cut between these shallow 
reef structures, but navigation through 
them is difficult and hazardous. Cover 
types range from unconsolidated with 
10% or less macroalgae cover to areas 
with greater than 10% coral or crustose 
coralline algae (NOAA, 2003; Figures 3.28, 3.29) 



57% 



o% 




■ Hardbottom with >10% live 
coral 

■ Hardbottom with >10% crustose 
coralline algae 

□ Hardbottom (uncolonized) 



35% 



□ Hardbottom with >10% 



macroalgae 

□ Hardbottom with indeterminate 
cover 

□ Unconsolidated with 10% or 
less macroalgae or seagrass 

■ Unconsolidated with >10% 
macroalgae or seagrass 



Figure 3.28. Percent composition of mapped benthic habitats at Maro Reef 
based on NOAA benthic habitat maps. Source: NOAA, 2003. 





BtnINc Habitat Type 



| HarfbMKm, HOHmoCrtiaJBB* [ | UnponiHniUlfl 



I Ufafddtud *#, * 10% ffl 
I * >„_„_ 



I 






L ^ \ 


Hab ita t Zones 

H Bartrtwf 
m DsOp rmrfeank 

.■■ ;■■■;: i '■ ■' N 
| Outer ruof ^ 
D a * 1 

iHZZi^iBK'torrKlcrs 









Figure 3.29. IKONOS satellite image (top left), benthic habitat map (top right), extent of water depth <20 m (bottom left), 
and habitat zones (bottom right) for Maro Reef. Maps: L. Wedding. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Because the outermost reefs absorb the majority of the energy from the open ocean swells, the innermost re- 
ticulated reefs and aggregated patch reefs are sheltered and have the characteristics of a true lagoon. Given 
the structural complexity of this platform, its shallow reefs are poorly charted and largely unexplored. 

Multibeam surveys were conducted at 
Maro Reef in 2002 (KM0206) and 2005 
(HI0508; Figure 3.30). The 2006 "ring" 
surveys on the perimeter were planned 
to delineate the 25-, 50-, and/or 100-fm 
isobaths for boundary designation pur- 
poses. 

At Maro, slope increases greatly be- 
tween the shallow (<226 m) and inter- 
mediate depths (226-588 m) and then 
declines slightly with greater depths 
(588-1,045 m). Aspect ratio declined 
slightly with depth from shallow to deep 
while rugosity was similar among all 
depths (Table 3.9). 



/ 



Bathymetry (20 m) 
Multibeam data 

depth (m) 



A 




Figure 3.30. 20 m Multibeam bathymetric data collected at Maro Reef in 
2002 and 2005 with derived depths from IKONOS imagery near island cen- 
ter. Map: L. Wedding. 



Table 3.9. Summary statistics for multibeam surveys conducted around Maro Reef (2002-2006, 


. 


MARO 


DEPTH CLASS 


AREA 


MINIMUM* 


MAXIMUM* 


RANGE 


MEAN 


STANDARD DEVIATION 


Bathymetry 


9 to 226 


900.29 


-259.40 


-1.00 


258.40 


-65.03 


56.96 


266 to 588 


277.74 


-616.72 


-202.22 


414.50 


-356.19 


106.15 


588 to 1,046 


133.73 


-800.00 


-559.44 


240.56 


-689.07 


61.27 


Slope 


9 to 226 


- 


0.00 


62.67 


62.67 


2.47 


4.30 


266 to 588 


- 


0.00 


64.64 


64.64 


8.48 


7.65 


588 to 1,046 


- 


0.00 


55.35 


55.35 


7.82 


5.76 


Aspect 


9 to 226 


- 


-1.00 


360.00 


361.00 


184.40 


108.58 


266 to 588 


- 


0.00 


360.00 


360.00 


175.34 


108.21 


588 to 1,046 


- 


-1.00 


360.00 


361.00 


155.68 


77.84 


Rugosity 


9 to 226 


- 


1.00 


2.23 


1.23 


1.00 


0.03 


266 to 588 


- 


1.00 


2.43 


1.43 


1.02 


0.04 


588 to 1,046 


- 


1.00 


2.08 


1.08 


1.02 


0.04 


*NOTE: the minimum represents the minimum value of a given 
mum value of a given metric within a universal depth class. 


metric within a 


jniversal depth class; max 


imum represents the maxi- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Laysan 

Laysan is a formed atoll, estimated to be 
20.7 million years old (Clague, 1996), 
with a maximum elevation of approxi- 
mately 15 m above sea level. By land 
area it is the second largest island in the 
Monument, with a land area of approxi- 
mately 3.7 km 2 , surrounded by close to 
405 km 2 of coral reef. Most of the reef 
area at Laysan lies in deeper waters, 
with a small, shallow water reef area in 
a bay off the southwest side of the is- 
land. It is well vegetated (except for its 
sand dunes) and contains a hyper-sa- 
line lake, which is one of only five natu- 
ral lakes in the state of Hawaii. Laysan's 
coral reef habitat totals approximately 
26.5 km 2 within 10 fathoms and 584.5 
km 2 out to 100 fathoms (Figures 3.31, 
3.32). The fringing reef surrounding the 
island varies from 100 to 500 m in width 



<i% 



5% <i% 



2% 




<1% 



■ Hardbottom with >10% live 
coral 

■ Hardbottom with >10% 
crustose coralline algae 

□ Hardbottom (uncolonized) 



□ Hardbottom with >10% 
macroalgae 

□ Hardbottom with indeterminate 
cover 

□ Unconsolidated with 10% or 
less macroalgae or seagrass 

■ Unconsolidated with >10% 
macroalgae or seagrass 



Figure 3.31. Percent composition of mapped benthic habitats at Laysan 
Island based on NOAA benthic habitat maps. Source: NOAA, 2003. 

and is most extensive at the northwest end of the island. Inside the 



171 4S-1Q-W 



17r*ttQTfY 




*&:Mi 




Benthic Hsbttat Type 



JHwdM 



i, >imkcnuniHO.LiraH™nlj» ^^ HwdOtfUm. 
vlQftnmciTMiaae 



I \J*itt*-K*iK*lw*h*-\Q%tr 



A 






■ 


Habitat Zones 

H[ E3.I' >. ■■::■;:' 

^ Qttp fftcr/bdrlk 

D 2 4 B 
^H=^^^H KiiMMtM 





Figure 3.32. IKONOS satellite image (top left), benthic habitat map (top right), extent of water depth <20 m (bottom left), 
and habitat zones (bottom right) for Laysan Island. Maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



reef is a narrow, shallow channel which 
nearly encircles the island except for 
the south and southeast sides. Deeper 
reef habitats are mostly robust spur- 
and-grooves around most of the island 
and a heavily eroded northern section 
with numerous caves, overhangs and 
large holes on a sloping reef with small 
sand channels. The base of these reefs 
and the spur-and-grooves ended in 
broad sand flats with numerous small 
overhangs and holes. Despite lacking 
much protection from the detrimental 
effects of waves, Laysan supports a 
surprisingly rich coral environment with 
good development along its leeward 
coasts. The small back reef, pass and 
moat near the island's western boat 
landing also help to diversify habitats 
and the number of coral species inhab- 
iting them. 



>»•»_, 



\ 




O 



4 



Bathymetry (20 m) 
Multibpam data 



\ Low: -1500 

^ikHnalars 



A 



Figure 3.33. 20-m multibeam data collected around Laysan Island (upper 
right) and Northampton Seamounts (lower left). Derived depths from IKO- 
NOS imagery are shown around Laysan Island. Map: L. Wedding. 



Multibeam surveys were conducted in 2002 during KM0206 in order to delineate 25, 50 and/or 100-fm bound- 
aries around Laysan Island (Figure 3.33). 



Lisianski-Neva Shoal 
Lisianski Island (Papaapoho) is another 
raised atoll, rising to 12.1 m above sea 
level, and with approximately 1.6 km 2 of 
emergent land is the third largest island 
within the Monument. This 23.4-million- 
year-old island (Clague, 1996) is over 
1.9 km across, consisting of an elevated 
rim surrounding a broad central depres- 
sion, although unlike Laysan it does not 
enclose an interior saline lake. The coral 
cover on the platform around the island, 
called Neva Shoal, is extensive, totaling 
over 1,174 km 2 (Figures 3.34, 3.35). 

Papaapoho describes a flat area with a 
depression or hollow, which is exactly 
how the island is shaped. Its highest 
point is a 12.2 m-high sand dune, and its 
lowest point is a depression to the south 
that runs as a channel toward the ocean. 



53% 




42% 



□ Hardbottom with >10% live 
coral 

■ Hardbottom with >10% 
crustose coralline algae 

□ Hardbottom (uncolonized) 



□ Hardbottom with >10% 
macroalgae 

□ Hardbottom with indeterminate 
cover 

□ Unconsolidated with 10% or 
less macroalgae or seagrass 

■ Unconsolidated with >10% 
macroalgae or seagrass 



Figure 3.34. Percent composition of mapped 
Neva Shoals based on NOAA benthic habitat 



benthic habitats at Lisianski- 
maps. Source: NOAA, 2003. 



Multibeam surveys were conducted in 2002 during KM0206 (Figure 3.36) in order to delineate 25, 50 and/or 
100-fm boundaries around Lisianski Island and Neva Shoals. The slope increases dramatically between the 
shallow (<226 m) and the intermediate depth range (226-588 m) before it declines slightly in the deeper depth 
bin (588-1,045 m). Aspect ratio is highest in the deep and shallow depth ranges while the intermediate depth 
ranges (266-588 m) had the lowest aspect ratio. Rugosity increased sharply between shallow and intermediate 
depths with a slight decrease in the deepest depth bin (Table 3.10). 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 











BoothiG Habttat Typ* 



□ 



CD 

13 



A 



iH'rw i7J"qw irysrw nvww 


tTJ'SffW 


fv 


HaWiet Zones 


^M 


i: .■;•>: reel 


^tI^^ ^1 ^^^ 


H DPOP W3*twnk 


^^^ ^j 


Lagoon roef 


^^fch ^1 lr 


1 Oulof r«f 




N 


^V^ V 7 






( JJ s to 





Figure 3.35. IKONOS satellite image (top left), benthic habitat map (top right), extent of water depth <20 m (bottom left), 
and habitat zones (bottom right) for Lisianski-Neva Shoals. Maps: L. Wedding. 




Bathymetry (20 m) 

Depth (m) 



2 4 8 



A 




Figure 3.36. 20 m multibeam data collected around Lisianski-Neva Shoal in 
2002 with derived depths from IKONOS imagery near island center. Map: 
L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 3.10. Summary statistics for multibeam 


surveys conducted around Lisianki (2002). 




LISIANKI 


DEPTH CLASS 


AREA 


MINIMUM* 


MAXIMUM* 


RANGE 


MEAN 


STANDARD DEVIATION 


Bathymetry 


9 to 226 


367.36 


-366.03 


-1.00 


365.03 


-47.61 


51.89 


266 to 588 


166.36 


-683.59 


-159.95 


523.64 


-430.03 


102.21 


588 to 1,046 


194.73 


-999.99 


-511.34 


488.65 


-785.34 


119.57 


Slope 


9 to 226 


- 


0.00 


78.38 


78.38 


4.34 


8.52 


266 to 588 


- 


0.00 


81.39 


81.39 


21.19 


12.56 


588 to 1,046 


- 


0.00 


72.36 


72.36 


19.10 


8.47 


Aspect 


9 to 226 


- 


-1.00 


360.00 


361.00 


174.50 


105.29 


266 to 588 


- 


-1.00 


360.00 


361.00 


149.92 


103.05 


588 to 1,046 


- 


-1.00 


360.00 


361.00 


177.09 


105.33 


Rugosity 


9 to 226 


- 


1.00 


6.83 


5.83 


1.02 


0.08 


266 to 588 


- 


1.00 


6.89 


5.89 


1.12 


0.19 


588 to 1,046 


- 


1.00 


3.39 


2.39 


1.08 


0.08 


*NOTE: the minimum represents the minimum value of a given metric within a universal depth class; maximum represents the maxi- 
mum value of a given metric within a universal depth class. 



Pearl and Hermes 

The name Holoikauaua celebrates the 
Hawaiian monk seals that haul out and 
rest here. Pearl and Hermes Atoll is a 
large atoll with several small islets, form- 
ing 0.38 km 2 of land surrounded by over 
1,214 km 2 of coral reef habitat (Figures 
3.37, 3.38). The atoll has an estimated 
age of 26.8 million years (Clague, 1996) 
and is over 32 km across and 19.3 km 
wide, with dunes rising above sea lev- 
el. Unlike Lisianski and Laysan to the 
southeast, Pearl and Hermes Atoll is a 
true atoll, fringed with shoals, permanent 
emergent islands, and ephemeral sandy 
islets. These features provide vital dry 
land for monk seals, green turtles, and 
a multitude of seabirds, with 16 species 
breeding here. The islets are periodi- 
cally washed over when winter storms 
pass through the area. 



5% 



-5% 




16% 



13% 



58% 



■ Hardbottom with >10% live 
coral 

■ Hardbottom with >10% 
crustose coralline algae 

□ Hardbottom (uncolonized) 



□ Hardbottom with >10% 
macroalgae 

□ Hardbottom with indeterminate 
cover 

□ Unconsolidated with 10% or 
less macroalgae or seagrass 

■ Unconsolidated with >10% 
macroalgae or seagrass 



Figure 3.37. Percent composition of mapped benthic habitats at Pearl and 
Hermes Atoll based on NOAA benthic habitat maps. Source: NOAA, 2003. 



Multibeam data were collected at Pearl and Hermes Atoll in 2005 (HI0509) to delineate the 25-, 50- and/ 
or 100-fm boundaries and in 2006 to complete as much mapping of the atoll as possible (Table 3.11; Figure 
3.39). Depths surveyed ranged from 9 m down to over 3,000 m. Slope increased by more than eight fold be- 
tween the shallowest (9-226 m) and the next deepest (226-588 m) depth ranges. The slope declined slightly 
in the deepest depth bin (1,649-3,071 m). Aspect ratio increased steadily with increasing depth, while rugosity 
rose sharply between shallow (<226 m) and intermediate ranges (226-1,046 m) and then declined slightly in 
the deepest depth bin (Table 3.11). Figure 3.40 shows a guyot approximately 25 km southeast of Pearl and 
Hermes Atoll where monk seals have been reported to forage through satellite tracking. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 







Figure 3.38. IKONOS satellite image (top left), benthic habitat map (top right), extent of water depth <20 m (bottom left), 
and habitat zones (bottom right) for Pearl and Hermes Atoll. Maps: L. Wedding. 

Table 3.11. Summary statistics for multibeam surveys conducted around Pearl and Hermes (2005) 



PEARL AND HERMES 


DEPTH CLASS 


AREA 


MINIMUM* 


MAXIMUM* 


RANGE 


MEAN 


STANDARD DEVIATION 




Bathymetry 


9 to 226 


611.14 


-1,818.20 


0.00 


1,818.20 


-44.22 


48.84 


266 to 588 


20.70 


-876.60 


-10.60 


866.00 


-336.52 


103.37 


588 to 1,046 


34.20 


-1,169.10 


-262.60 


906.50 


-862.04 


123.81 


1,046 to 1,649 


92.48 


-1,858.90 


-631.80 


1,227.10 


-1,368.91 


175.49 


1,649 to 3,071 


86.96 


-3,071.30 


-40.70 


3,030.60 


-1,979.21 


281.68 


Slope 


9 to 226 


- 


0.00 


77.41 


77.41 


3.39 


4.73 


266 to 588 


- 


0.00 


82.89 


82.89 


28.25 


11.14 


588 to 1,046 


- 


0.00 


84.42 


84.42 


28.60 


11.14 


1,046 to 1,649 


- 


0.00 


84.72 


84.72 


22.32 


10.32 


1,649 to 3,071 


- 


0.00 


87.49 


87.49 


15.04 


9.89 


Aspect 


9 to 226 


- 


-1.00 


360.00 


361.00 


158.44 


116.76 


266 to 588 


- 


-1.00 


359.94 


360.94 


198.48 


97.80 


588 to 1,046 


- 


-1.00 


360.00 


361.00 


202.81 


95.63 


1,046 to 1,649 


- 


-1.00 


360.00 


361.00 


224.08 


98.92 


1,649 to 3,071 


- 


-1.00 


360.00 


361.00 


225.85 


98.48 


Rugosity 


9 to 226 


- 


1.00 


14.14 


13.14 


1.01 


0.03 


266 to 588 


- 


1.00 


12.93 


11.93 


1.30 


0.43 


588 to 1,046 


- 


1.00 


18.72 


17.72 


1.32 


0.48 


1,046 to 1,649 


- 


1.00 


19.01 


18.01 


1.25 


0.38 


1,649 to 3,071 


- 


1.00 


89.27 


88.27 


1.16 


0.46 


*NOTE: minimum = minimum 


/alue of a given metric within a un 


iversal depth 


class; maximun 


l = the maximur 


n value of a given metric wit 


lin a universal depth class. 






A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Bathymalry (5 m) 
MulEl beam data 

Oepm (m) 
■■ o 

N 

a 1 i a ™ 



175TKW ITS'H'W CHP0W 17F«Tff 




175"S4°W ITSMflW 



HS'HW IK'ttW 



Ru-goslty (SmJ 
MuJlibeam da la 




1TS"54TW 175"«'W 



N 

1 2 * ... " 



lWQTff whin 




l'fl"W 1W5*"W 



17IWW 17S , H'W 




ITfi-eiff 17&-IVW 



175"5*W 175"«'W nffyawi 



Figure 3.39. 5 m and 20 m bathymetry, rugosity (5 m), and slope (5 m) for Pearl and Hermes with derived depths from 
IKONOS imagery near island center. Maps: L. Wedding. 



1/S39W IJbJE'W 



UblJW 



Seamount SE of Pearl & Hermes Atoll, MWHI 
NOM Ship Hiialakai Multibeam Balhymelry 
40 m grid cell size - C.I. = 50 m 




■za» -sot -is» -ifflo -nm -1200 -now -m -wo ■*« -2w 
BWUVMETRY (m) 




Figure 3.40. 3-D perspective of a seamount southeast of Pearl and Hermes Atoll. V.E. = 2x. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Midway Atoll 

Midway Atoll consists of three sandy is- 
lets: Sand, 4.56 km 2 ; Eastern, 1.36 km 2 ; 
and Spit, 0.05 km 2 for a total of 1,464 
5.9 km 2 in terrestrial area, lying within a 
large, elliptical barrier reef measuring ap- 
proximately 8 km in diameter. The atoll, 
which is 28.7 million years old (Clague, 
1996), is surrounded by more than 356 
km 2 of coral reefs (Figures 3.41, 3.42). In 
1965, the U.S. Geological Survey took 
core samples and hit solid basaltic rock 
54.8 m beneath Sand Island and 377.9 
m beneath the northern reef. Numerous 
patch reefs dot the sandy-bottomed la- 
goon. The atoll and surrounding seas 
were also the site of a pivotal battle of 
World War II, and Midway was an active 
Navy installation during the Cold War. 



<i%- 



i% 



-<i% 



53% 




IHardbottom with >10% live coral 



■ Hardbottom with >10% crustose 
coralline algae 

□ Hardbottom (uncolonized) 



□ Hardbottom with >10% 
macroalgae 

□ Hardbottom with indeterminate 
23% cover 

□ Unconsolidated with 10% or less 
macroalgae or seagrass 

■ Unconsolidated with >10% 
macroalgae or seagrass 



7% 



Figure 3.41. Percent composition of mapped benthic habitats at Midway 
Atoll based on NOAA benthic habitat maps. Source: NOAA, 2003. 



Multibeam mapping surveys at Midway were conducted in 2003 (AHI0306) to delineate 25-, 50- and 100-fm 
boundaries and in 2005 (HI0503) and 2006 (HI0609) to add to coverage around the island for benthic habitat 



1T-Z7W 




177 24-W 


17T-31-W 












1 | 














k 




i 








1 




V 


H&£ v 


iflm 


HHHJS 


V 


H 




-~M— — 


^ 






A °_r 


1 2 













Benthic H&bitat Type 



IJn«i 



1 Ufn»HK*inl»d wrttt "= 1 OK rr 



A 



Hibifetzon** 




^| R.V i!l-l 




^H D**D rWMsnil 




Lftg«Kir** 




[ | Oitinr n»r 




"71 Land 


N 

A 


Vfate <2fl m 


& 1 2 4 




Figure 3.42. IKONOS satellite image (top left), benthic habitat map (top right), extent of water depth <20 m (bottom left) 
and habitat zones (bottom right) for Midway Atoll. Maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

mapping (Figure 3.43). Slope increased by 10 fold between the shallowest and intermediate depth with a mod- 
est decline in the deepest depth bin. Aspect ratio increased moderately between shallow and intermediate 
depth but declined sharply in the 588-1,045 m depth range. Rugosity increased from shallow to intermediate 
before declining slightly in the deepest depth range (Table 3.12). 











s 



\ 



Ryywtty (5 ■■■) 
Mull ■.«' '.\i> dais 



, A 




Bathymetry (20 m) 
Multl beam data 




Figure 3.43. 5 m and 20 m bathymetry, rugosity (5 m), and slope (5 m) for Midway Atoll with derived depths from IKONOS 
imagery near island center. Maps: L. Wedding. 

Table 3.12. Summary statistics for multibeam surveys conducted around Midway Atoll (2003-2006). 




MIDWAY 


DEPTH CLASS 


AREA 


MINIMUM* 


MAXIMUM* 


RANGE 


MEAN 


STANDARD DEVIATION 


Bathymetry 


9 to 226 


306.19 


-250.60 


3.00 


253.60 


-60.98 


51.15 


266 to 588 


37.32 


-500.00 


2.00 


502.00 


-349.27 


77.63 


588 to 1,046 


0.00 


-2.00 


-2.00 


0.00 


-2.00 


0.00 


Slope 


9 to 226 


- 


0.00 


56.57 


56.57 


2.85 


5.16 


266 to 588 


- 


0.00 


79.34 


79.34 


28.83 


9.04 


588 to 1,046 


- 


19.97 


19.97 


0.00 


19.97 


0.00 


Aspect 


9 to 226 


- 


-1.00 


360.00 


361.00 


137.68 


114.67 


266 to 588 


- 


-1.00 


360.00 


361.00 


170.08 


95.52 


588 to 1,046 


- 


40.82 


40.82 


0.00 


40.82 


0.00 


Rugosity 


9 to 226 


- 


1.00 


2.29 


1.29 


1.01 


0.04 


266 to 588 


- 


1.00 


4.87 


3.87 


1.19 


0.16 


588 to 1,046 


- 


1.11 


1.11 


0.00 


1.11 


0.00 


*NOTE: the minimum represents the minimum value of a given metric within a universal depth class; maximum represents the maximum value of a 
given metric within a universal depth class. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Ku re Atoll 

Kure Atoll is the most northwestern island in the Hawaiian chain and occupies a singular position at the "Darwin 
Point": the northern extent of coral reef development, beyond which coral growth cannot keep pace with the 
rate of geological subsidence. Kure's coral is still growing slightly faster than the island is subsiding. North of 
Kure, where growth rates are even slower, the drowned Emperor Seamounts foretell the future of Kure and all 
of the Hawaiian Archipelago. As Kure Atoll continues its slow migration atop the Pacific Plate, it too will eventu- 
ally slip below the surface (Grigg, 1982). 



This 29.8 million year old atoll (Clague, 
1996) is nearly circular, with a reef 9.6 
km in diameter enclosing a lagoon with 
two islets that include over 0.81 km 2 of 
emergent land, flanked by almost 324 
km 2 of coral reef habitat (Figures 3.44, 
3.45). The outer reef forms a nearly 
complete circular barrier around the la- 
goon, with the exception of passages 
to the southwest. Of the two enclosed 
islets, the only permanent land is found 
on crescent-shaped Green Island, 
which rises to 6.1 m above sea level and 
is located near the fringing reef in the 
southeastern quadrant of the lagoon. 
The USCG established a LORAN sta- 
tion at Kure in 1960 (Woodward, 1972) 
and occupied it until 1993. This land use 
had far-reaching effects on all the plants 
and animals at Kure Atoll, resulting in 
elevated invasive species problems and 
contaminants left behind when the base 
closed. 





4% x 


3% 


1% 


v 17% 




■ Hardbottom with >10% live coral 

■ Hardbottom with >10% crustose 
coralline algae 

□ Hardbottom (uncolonized) 














8% 


□ Hardbottom with >10% 
macroalgae 














□ Hardbottom with indeterminate 
cover 


55% \ 








/\ 

/ 3 




□ Unconsolidated with 10% or less 
macroalgae or seagrass 










L2% 


■ Unconsolidated with >10% 
macroalgae or seagrass 











Figure 3.44. Percent composition of mapped benthic habitats at Kure Atoll 
based on NOAA benthic habitat maps. Source: NOAA, 2003. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



17S 2AW 1 78-21 TAI 


ira-ifl-w 






1 > 


it^k 




4t* 




fe 


J 


17H + 24W 17^21 Vf 


i.'i! ; ;w 




I urhaynonhod with < Igtt » wri t« 
N 

° " ■ '■■■■ A 



Hw 





A 


1 !. . :: i-. .: Zww 
■1 Bacfciwf 
|^| De*p war.tMink 

Qutarraar i^j 
15 3 8 


^^% 









Figure 3.45. IKONOS satellite image (top left), benthic habitat map (top right), extent of water depth <20 m (bottom left), 
and habitat zones (bottom right) for Kure Atoll. Maps: L. Wedding. 

Multibeam data were collected around Kure Atoll in 2005 (HI0509) to delineate the 25, 50 and 100-fm bound- 
aries and in 2006 (HI0609) to complete mapping of the atoll (Figure 3.46). Slope increased by 9 fold between 
the shallow and intermediate depth ranges (Table 3.13). Aspect ratio and rugosity also showed the same trend 
although the magnitude was not as great (Figure 3.47). Multibeam bathymetry at Kure and Midway show ex- 
tensive spur and groove formations in the high resolution data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



BallijiiitLrir f5 in} 

Mull.b 

Dnpl.ii 



1.252.5 5 






y \ 




Rugosity (5 m> 
Hlin beam data 


J 


J 

I 


Rupasity 

ma Hqh 




xh 


[ :-v 

N 

1 252 5 5 ^ 

^■B=HHH KilwTMlfhrs 




J 









Bathymetry (20 m) 
Doer 




Slope fSm) 
MuHl beam data 


Slope 




^B vnpstanp 


S5" 


VofyFkitti 


N 

A 


L.|n,J 

D 1.252.5 5 




Figure 3.46. 5 m and 20 m bathymetry, rugosity (5 m), and slope (5 m) for Kure Atoll. Maps: L. Wedding. 



Table 3.13. Summary statistics for multibeam surveys conducted around Kure Atoll (2005-2006). 


KURE 


DEPTH CLASS 


AREA 


MINIMUM* 


MAXIMUM* 


RANGE 


MEAN 


STANDARD DEVIATION 


Bathymetry 


9 to 226 


323.34 


-225.00 


1.00 


226.00 


-73.27 


52.46 


266 to 588 


30.76 


-402.00 


-226.00 


176.00 


-314.77 


50.63 


Slope 


9 to 226 


- 


0.00 


77.44 


77.44 


2.86 


4.68 


266 to 588 


- 


0.00 


77.82 


77.82 


26.69 


8.43 


Aspect 


9 to 226 


- 


-1.00 


359.43 


360.43 


153.84 


117.63 


266 to 588 


- 


-1.00 


359.79 


360.79 


183.77 


97.63 


Rugosity 


9 to 226 


- 


1.00 


7.63 


6.63 


1.01 


0.03 


266 to 588 


- 


1.00 


8.12 


7.12 


1.15 


0.12 


*NOTE: the minimum represents the minimum value of a given metric within a universal depth class; ma 
mum value of a given metric within a universal depth class. 


ximum represents the maxi- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Figure 3.47. Spur and groove formations show up clearly in the high resolution bathymetry taken around Kure Atoll. 




sa 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



EXISTING DATA GAPS 



• A comprehensive shallow- water benthic habitat map is required to support multiple research and monitor- 
ing activities. Currently only about 50% of the shallow-water area from 0-30 m has sufficient coverage and 
quality of imagery to produce maps. 

• A complete and seamless digital terrain and bathymetric models for the Papahanaumokuakea Marine 
National Monument (PMNM) to at least 500 m is currently lacking and is necessary to better understand 
the connection of shallow and deep-water habitats. It will also help guide assessment and monitoring 
activities. 

• Need to complete high-resolution multibeam bathymetry for shallow to moderate (20 to 500 m) 
depths. 

• Need to collect shallow (<20 m) bathymetric data using Light Detection and Ranging (LIDAR) or other 
appropriate technologies. 

• Sea level curves developed from the MHI and elsewhere around the Pacific might not be appropriate 
for the NWHI. Fossil coring and other methods should be employed to better understand past sea level 
changes in the NWHI. 

• Need to determine whole reef accretion rates in different geomorphological zones. 

• Need to examine rates of terrestrial habitat loss and the factors that cause it. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

REFERENCES 

Clague, D.A. 1996. The growth and subsidence of the Hawaiian-Emperor volcanic chain. Pp: 35-50. In: Keast A., S.D. 
Miller (eds.). The origin and evolution of the Pacific Island biota, New Guinea to Eastern Polynesia: patterns and pro- 
cesses. SPB Academic Publishing, Amsterdam, The Neatherlands. 

Clague, D.A. and G.B. Dalrymple. 1987. The Hawaiian-Emperor volcanic chain: Part I. Geologic evolution. US Geological 
Survey Professional Paper 1350, pp. 5-54. 

Clapp, R.B. 1972. The natural history of Gardner Pinnacles, northwestern Hawaiian Islands: botany. Atoll Res. Bull., 
no. 163. 

Davies, T.A., P. Wilde, and D.A. Clague. 1972. Koko seamount: a major guyot at the southern end of the Emperor sea- 
mounts. Mar. Geol. 13: 311-321. 

Garcia, M., D. Grooms, and J. Naughton. 1987. Petrology and geochronology of volcanic rocks from seamounts along 
and near the Hawaiian Ridge. Lithos 20: 323-336. 

Grigg, R. W. 1981. Coral reef development at high latitudes in Hawaii, Proceedings of the Fourth International Coral Reef 
Symposium, Manila, 1, 687-693. 

Grigg, R.W. 1982. Darwin Point: a threshold for atoll formation. Coral Reefs 1: 29-34. 

Grigg, R.W. 1983. Community structure, succession and development of coral reefs in Hawaii. Mar Ecol Prog Ser 11:1- 
14. 

Grigg, R.W. 1997. Paleoceanography of coral reefs in the Hawaiian-Emperor Chain - Revisited. Coral Reefs 16, 
Suppl:S33-S38. 

Grigg, R.W., J. Polovina, A. Friedlander, and S. Rohmann. 2008. Pp: 573-594 in: Biology and paleoceanography of the 
coral reefs in the northwestern Hawaiian Islands. In: Coral reefs of the United States (B. Riegl and R. Dodge, eds.). 
Springer-Vergal Publishing. 

Gross M.G., J.D. Milliman, J.I. Tracey, and H.S. Ladd. 1969. Marine geology of Kure and Midway Atolls, Hawaii: a pre- 
liminary report. Pac. Sci. 23:17-25. 

Hoeke, R., R. Brainard, R. Moffitt, and M. Merrifield. 2006. The role of oceanographic conditions and reef morphology in 
the 2002 coral bleaching event in the northwestern Hawaiian Islands. Atoll Res. Bull. 543:489-503. 

Jackson, E. D.; Shaw, H. R.; Bargar, K. E. Calculated geochronology and stress field orientations along the Hawaiian 
chain. Earth and Planetary Science Letters, Vol. 26, p. 145 

Jenness, J. 2003. Grid surface areas: surface area and ratios from elevation grids [electronic manual]. URL http://www. 
jennessent.com/arcview/arcview_extensions.htm 

Juvik, S.P and J.O. Juvik. 1998. Atlas of Hawaii, 3rd edition. University of Hawaii Press. Honolulu. 

Lundblad, E., D.J. Wright, J. Miller, E.M. Larkin, R.B. Rinehart, S.M. Anderson, D.F. Naar, and B.T Donahue. 2006. A 
benthic terrain classification scheme for American Samoa. Marine Geodesy 29: 89-111. 

Maragos, J.E., D.C. Potts, F Aeby, D. Gulko, J. Kenyon, D. Siciliano, and D. VanRavenswaay. 2004. 2000-2002 Rapid 
Ecological Assessment of Corals (Anthozoa) on Shallow Reefs of the Northwestern Hawaiian Islands. Part 1. Species and 
Distribution. Pacific Science 58(2): 211-230. 

Miller, J.E., R.K. Hoeke, T.B. Appelgate, P.J. Johnson, J.R. Smith, and S. Bevacqua. 2003. Atlas of the Northwestern Ha- 
waiian Islands, Draft - February 2004, National Oceanic and Atmospheric Administration, 65 pp. 

Neall, V.E. and S.A. Trewick. 2008. The age and origin of the Pacific islands: a geological overview. Phil. Trans. R. Soc. 
B 363:3293-3308. 

NOAA (National Oceanic and Atmospheric Administration). 2003. Atlas of the Shallow-Water Benthic Habitats of the 
Northwestern Hawaiian Islands (Draft). 160 pp. 

Parrish, FA. and J.J. Polovina. 1994. Habitat thresholds and bottlenecks in production of the spiny lobster (Panulirus 
marginatus) in the northwestern Hawaiian Islands. Bulletin of marine science 54: 151-163. 

Rohmann, S.O., J.J. Hayes, R.C. Newhall, M.E. Monaco, and R.W. Grigg. 2005. The area of potential shallow-water tropi- 
cal and subtropical coral ecosystems in the United States. Coral Reefs 24:370-383. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Rooney, J.J., P. Wessel, R. Hoeke, J. Weiss, J. Baker, F. Parrish, C.H. Fletcher, J. Chojnacki, M. Garcia, R. Brainard, and 
P. Vroom. 2008. Geology and Geomorphology of Coral Reefs in the Northwestern Hawaiian Islands. In: Coral reefs of the 
United States (B. Riegl and R. Dodge, eds.). Springer-Vergal Publishing. 

Stumpf, R.P. and K. Holderied. 2003. Determination of water depth with high-resolution satellite imagery over variable 
bottom types. Limnol. Oceanogr. 48: 547-556. 

Stumpf, R.P, K. Holderied, and M. Sinclair. 2003. Determination of water depth with high-resolution satellite imagery over 
variable bottom types. Limnology and Oceanography 48: 1. 

Woodward, PW. 1972. The natural history of Kure Atoll, Northwestern Hawaiian Islands. Atoll Res. Bull. 164:1-317. 

WEBSITES 

NOAA. 2008. http://ccma.nos.noaa.gov/ecosystems/coralreef/nwhi/welcome.html 
NOAA. 2008. http://biogeo.nos.noaa.gov 

UH SOEST. 2008. http://www.soest.hawaii.edu/pibhmc/pibhmc_nwhi.htm 
NOAA. 2008. http://biogeo.nos.noaa.gov 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Benthic Communities 

James Maragos 1 , Jean Kenyon 23 , Greta Aeby 4 , Peter Vroom 23 , Bernardo Vargas-Angel 23 , Russell Brainard 2 , 
Lisa Wedding 45 , Alan Friedlander 56 , Jacob Asher 23 , Brian Zgliczynski 2 and Daria Siciliano 7 

INTRODUCTION 

This chapter focuses on the shallow water benthos surrounding the 10 emergent islands, reefs and atolls 
comprising the Northwestern Hawaiian Islands (NWHI). The composition and distribution of benthic commu- 
nities in the subtropical NWHI reflect the interaction of numerous influences, including geographic isolation, 
latitude, exposure and the successional age of the islands they inhabit. This mosaic of habitats extend 1,200 
nm northwest of the main Hawaiian Islands (MHI) and consist mostly of coral-dominated areas, stretches of 
hard-bottom algal-dominated meadows, and vast expanses of unconsolidated sediments such as sand and 
mud inhabited by few benthic infauna at shallower depths (<20 m). The benthic habitats at greater depths are 
relatively unknown, unexplored and are not covered here. 



CORALS 

Species Richness 

The most recent published surveys of the NWHI revealed a total of 57 species of zooxanthellate stony corals 
(Maragos et al., 2004). However 2006 surveys at greater depths and in a wider variety of habitats have yielded 
a number of new morpho-species, most of which have yet to be collected and described. Appendix I lists of all 
coral and anemone species reported at 11 islands, banks, atolls and reefs in the NWHI as of October 2006. As 
was the case during earlier compilations, 
the larger atolls with diverse habitats 
and shelter from large northwest swell 
support the greatest number of species. 
Although these numbers are similar to 
the MHI, the Hawaiian coral fauna, as 
a whole, is depauperate relative to the 
Indo-West-Pacific, where up to 700 spe- 
cies have been reported (Veron, 1995). 
The most plausible cause is geographic 
isolation (Grigg, 1983) associated with 
the NWHI being located at the north- 
eastern periphery of the Indo-Pacific 
biogeographic province. 

The distribution of coral species is re- 
lated to geomorphology, size and age 
of the NWHI reefs (Grigg, 1997; Figure 
4.1). There is a highly significant (p< 
0.001) correlation between the number 
of coral taxa and the amount of reef 
area within 10 fathoms with French Frig- 
ate Shoals, Pearl and Hermes and Maro 




Figure 4.1. Total number of coral species reported in the NWHI between 
1907 and 2006 compiled by Maragos from Dana (1846), Vaughan (1907), 
Dana (1971), Maragos et al. (2004) and unpublished records. Map: L. Wed- 
ding. 



1. U.S. Fish and Wildlife Service 

2. NOAA/NMFS/Pacific Islands Fisheries Science Center, Coral Reef Ecosystem Division 

3. Joint Institute for Marine and Atmospheric Research 

4. University of Hawaii at Manoa 

5. NOAA/NOS/NCCOS/CCMA Biogeography Branch 

6. The Oceanic Institute 

7. University of California Santa Cruz, Institute of Marine Sciences 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Reef having the highest richness of cor- 
al taxa (Table 4.1, Figure 4.2). Low coral 
species richness is found at the south- 
ern end among the small basalt islands 
of Nihoa, Mokumanamana and Gardner 
Pinnacles, which are openly exposed to 
severe wave events particularly during 
winter months (Maragos et al., 2004 ; 
Grigg et al., 2008). This area is dominat- 
ed by robust species that are more tol- 
erant of high wave action and soft cor- 
als such as Sinularia spp. and Palythoa 
spp. The large middle atolls (French 
Frigate Shoals and Maro) have some 
of the highest coral species richness 
reflecting optimal conditions in terms 
of both habitat (a large open atoll) and 
environmental conditions (wave shelter, 
temperature and low disturbance, as 
well as the high number of Acroporidae 
and Fungiidae species). Seven species 
of the genus Acropora are now known 
from the NWHI despite their almost com- 
plete absence in the MHI (Maragos et 
al., 2004), and several additional unde- 
scribed Acropora were photographed at 
French Frigate Shoals, Neva Shoal and 
Pearl and Hermes in 2006. Compared 
to French Frigate Shoals, Pearl and 
Hermes Atoll and Maro Reef, one-third 
fewer coral species are found at Moku- 
manamana and Nihoa islands. At the 
northern end of the chain, stony coral 
species decline is linked to lower winter 
water temperatures and lower average 
annual solar radiation (Grigg, 1982), and 
coral development is limited by exten- 
sive sand shallows towards the eastern 
sides of the lagoons (Maragos et al., 
2004). Moreover WWII era construction 
may have extirpated some corals from 



Table 4.1. Least squares linear regression model for reef area less than 
10 fathoms versus coral taxa richness. Source: Maragos, unpub. data. 



SUMMARY OF FIT 


R 2 


0.79859 


R 2 Adjusted 


0.773414 


Root Mean 
Square Error 


6.390495 


Mean of Response 


35.7 


Observations 
(or Sum Wgts) 


10 


ANALYSIS OF VARIANCE 








Source 


DF 


Sum of Squares 


Mean Square 


F Ratio 


Model 


1 


1295.393 


1295.39 


31.7199 


Error 


8 


326.7074 


40.84 


Prob > F 


C. Total 


9 


1622.1 




0.0005 


PARAMETER ESTIMATES 








Term 


Estimate 


Std Error 


t Ratio 


Prot»|t| 


Intercept 


24.88415 


2.7878 


8.93 


<.0001 


Reef area 
< 10 fathoms 


0.072376 


0.012851 


5.63 


0.0005 



100 



80 



60 



O 

H 40 



20 



Reef area < 10 fathom vs coral species richness 
Linear regression line 
95% Confidence Band 
95% Prediction Band 




100 



200 



300 



400 



500 



Reef area <10 fathoms (km ) 



Figure 4.2. Relationship between number of coral taxa and reef area within 
10 fathoms (based on Rohman et al., 2005). Coral species richness = 24.88 
+ 0.072*Reef area <10 fathoms. Source: Maragos, unpub. data. 



Midway's lagoon. Kure is the world's most northern atoll and is referred to as the Darwin Point, where coral 
growth and subsidence/erosion balance one another (Grigg, 1982). However, there are at least 30 submerged 
and presumably drowning seamounts to the southeast of Kure and within the PMNM, and the "Darwin point" 
may be better characterized as a Darwin zone stretching from Nihoa to Kure. 

Examination of reefs in ordination space based on presence-absence of coral taxa reveal two major clusters 
(Figure 4.3). High concordance exists among the basalt islands of Nihoa (NIH), Mokumanamana (MMM) and 
Gardner Pinnacles (GAR). These small islands are exposed to high wave energy from all directions and have 
low coral richness and cover. The reefs from Maro (MAR) north to Kure (KUR) cluster together in ordination 
space but the three most northern atolls (Midway [MID], Pearl and Hermes [PHR], and Kure [KUR]) show the 
highest concordance. The coral assemblage at French Frigate Shoals appears unique compared with all other 
locations likely due to the high proportion of acroporid species and possible connectivity with Johnston Atoll 
830 km to the south (Grigg et al., 1981; Maragos and Jokiel, 1986). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Range Extensions and Possible New 
Coral Species 

Recent scientific expeditions in the 
NWHI have yielded many new records 
and possibly many undescribed species 
of stony corals since the last compilation 
by Maragos et al. (2004; Table 4.2). One 
of the most exciting discoveries in 2006 
was of the table coral Acropora off the 
southwest spur-and-groove habitat at 
Pearl and Hermes Atoll and off the shal- 
low southeast fore reef at Neva Shoal. 
Additional dives confirmed the pres- 
ence of Acropora cytherea and A. cere- 
alis-valida at Pearl and Hermes, and A. 
valida at Neva Shoal, which led to other 
discoveries at Neva, a second Acropora 
sp.l and three Montipora species that 
are all likely new to science. 




Figure 4.3. Spatial distribution of reefs in the PMNM based on presence- 
absence of coral species. Results of non-metric multidimensional scaling 
plot. Source: Maragos, unpub. data. 



The Census of Marine Life (CoML) cruise to French Frigate Shoals in October 2006, led to additional sightings 
of rare species including Diaseris distorta, Cycloseris tenuis, Leptoseris scabra and Acropora sp.l. Another 
rare species, resembling Leptoseris papyracea was previously known only from dredge hauls by Vaughan 
(1907) in the MHI, and was reported for the first time in the NWHI off the southeast fore reef of French Frigate 
Shoals during CoML. An unidentified species, Pontes sp. 15, was reported off a southwest pinnacle of French 
Frigate Shoals, and the first record of Porites lutea in the NWHI was reported off the northern reef crest. Many 
new records including several unidentified species of coral were reported during the CoML cruise. The 2006 
investigations together have yielded possibly 11 new records for the NWHI most of which are likely to reveal 
new species. Towed-diver surveys contributed directly or indirectly to several of the new records and species, 
and exploratory dives in new habitats and sites contributed the rest. The Appendix at the end of the chapter 
lists all coral and anemone species reported at 11 islands, banks, atolls and reefs in NWHI as of October 2006. 
As was the case during earlier compilations, the larger atolls with diverse habitat and shelter from large north- 
west swell support the greatest number of species. 



Table 4.2. A partial list of new records and possible new coral species from the NWHI. Source: Maragos, unpub. data. 


SPECIES 




LOCATION 


YEAR 


OBSERVERS 


Leptoseris incrustans 


New record 


Pearl and Hermes 


2006 


Kenyon, PIFSC-CRED 


Montipora 


New species? 


Pearl and Hermes 


2006 


Vargas, PIFSC-CRED 


Acropora valida 


New record 


Laysan 


2006 


Kenyon, PIFSC-CRED 


Pavona maldivensis 


New record 


Maro 


2006 


Kenyon, PIFSC-CRED 


Diaseris distorta, Cycloseris vaughani, and Cycloseris tenuis and soft 
coral Sinularia sp 


New record 


Lisianski (30m) 


2006 


Maragos, Meyer, and 
Papastamatiou 


Acropora cytherea and A. cerealis-valida 


New record 


Pearl and Hermes 


2006 


Asher and Zgliczynski 


Acropora valida 


New record 


Neva Shoal 


2006 


Asher and Zgliczynski 


Diaseris distorta, Cycloseris tenuis, Leptastrea scabra 


New record 


French Frigate Shoals 


2006 


CoML 


Leptoseris papyracea?? 


New record 


French Frigate Shoals 


2006 


CoML 


Acropora sp.l 


New species? 


French Frigate Shoals 


2006 


CoML 


Porites sp. 15 


New species? 


French Frigate Shoals 


2006 


CoML 


Porites lutea 


New record 


French Frigate Shoals 


2006 


CoML 


Unkown species 


New species? 


French Frigate Shoals 


2006 


CoML 


Abbreviations: PIFSC-CRED: The Pacific Islands Fisheries Science Center Coral Reef E 


:osystem Division; CoML 


.=Censu 


s of Marine Life. 





Figure 4.4. Potential new species of Acropora sp.l from Lisianski-Neva 
Shoals and French Frigate Shoals (left). New, potential relict species yet 
undescribed from French Frigate Shoals. The genus and family are un- 
known (right). Both were discovered in 2006. Photo: J. Maragos. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

The most exciting coral discovery was 
of an unknown species that has not yet 
be identified to the genus and family 
(Figure 4.4). This coral may be a rel- 
ict that was once common in the past 
and subsequently died out elsewhere 
but survived in Hawaii. The other pos- 
sibility is that the coral may be a type 
previously restricted to deep water that 
evolved and subsequently adapted it- 
self to shallow water habitats. Randall 
(2007) makes note of two fish that were 
previously characterized as relicts. Like- 
wise it is possible for relict corals to have 

survived in Hawaii to this day. In order to confirm this, it will be necessary to collect this and other corals to 
determine their phylogenetic origin. So far coral experts have not been able to conclusively determine the fam- 
ily to which this coral belongs based on photographs alone. Marine life in the NWHI evolved for many millions 
of years in isolation from neighboring archipelagos and islands and it plausible that this, and perhaps other 
species, were able to survive and thrive without the threat of newer species displacing them as likely occurred 
in other archipelagos. 

French Frigate Shoals was chosen as the target for the first CoML cruise because of the potential of yielding 
new species of corals, other invertebrates and benthic algae and possibly extending the range of many other 
species. Eight more species of cnidarians have already been reported from the atoll, further cementing the 
atoll's status as the most diverse island or atoll for corals in Hawaii. The atoll is the closest of the Hawaiian 
chain to Johnston Atoll, some 450 nm to the southwest, and Johnston may be serving as a "stepping stone" for 
the dispersal of species to Hawaii from the Line Islands and other neighboring archipelagos south of Hawaii 
(Grigg, 1981; Maragos and Jokiel, 1986; Maragos et al,. 2004). This connection would explain why French 
Frigate Shoals has so many Acropora species which flourish at Johnston and why French Frigate Shoals has 
higher numbers of coral species compared to any of the other Hawaiian Islands. 



Coral Endemism 

The resumption of coral surveys to the 
NWHI in 2000-2006 were not focused on 
looking for new coral species because 
they were not expected to be there 
based on the Grigg and Dollar (1980) 
estimate of only 29 total species at 80 
NWHI sites. Although endemic species 
and range extensions were encountered 
during the 2000-2006 surveys, most be- 
longed to described species, although 
the totals nearly doubled the number 
of coral species (57) reported in the 
NWHI (Maragos et al., 2004). The ex- 
ploratory and CoML expeditions of 2006 
were both focused on looking for new 
species. However, scientists could only 
photograph and examine corals in situ 
because permission was not granted to 
collect corals. 




Figure 4.5. Percent endemic stony coral species at each reef in the NWHI 
during 1907-2006 compiled by Maragos from Dana (1846), Vaughan 
(1907), Dana (1971), Maragos et al. (2004) and unpublished records. Map: 
L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Based on all surveys to date (Figure 
4.5; Table 4.3) , there are now approxi- 
mately 80 morphologically distinct coral 
"species" in the NWHI, and about 35 are 
likely to be endemic. Perhaps an addi- 
tional five species are similarly endemic, 
although other specialists (Veron, 2000) 
believe they are widespread Pacific 
species. More than 25 NWHI species 
are still undescribed or unidentified, and 
once type specimens are collected and 
examined morphologically and geneti- 
cally, then final determinations can be 
made on which corals are new species 
and possible endemics. Notwithstand- 
ing efforts to date, NWHI explorations 
are still inadequate, and it is likely that 
additional undescribed coral species 
will be encountered in the future. 



Coral Cover From Quantitative 
Surveys 

Percent live coral cover was derived 
from towed-diver data and rapid ecolog- 
ical assessments (REAs; Kenyon et al,. 
2006a; Table 4.4; Figures 4.6 and 4.7). 
Towed-divers observe a much greater 
expanse of benthic reef habitat (approx- 
imately 2 km in length/50 minute tow) 
than can be observed by free-swim- 
ming divers conducting REA surveys 
in a comparable time period and give a 
more widespread assessment of the to- 
tal coral cover for each reef. Coral REA 
surveys were conducted at 70 sites in 
2004, 37 sites in 2005 and 64 sites in 
2006 (Table 4.4). As with percent cover 
data from 2002 surveys (which were 
calculated from size frequency data of 
colony counts within transects; Fried- 
lander et al.; 2005), line-intercept data 
from surveys in all three years indicated 
coral cover varies greatly across the 
NWHI. 

Based on towed-diver survey data, 
Lisianski (18.8%) and Maro (14.9%) 
have the highest coral cover, followed 
by French Frigate Shoals (Figure 4.6). 
Percent coral cover from the REAs is 
higher owing to the fact that they were 
conducted on hard bottom habitats only. 
However, the trends are strikingly simi- 



Table 4.3. Number of endemic and total stony coral taxa among major reefs 
in the PMNM. Source: Maragos, unpub. data. 


ISLAND 


TOTAL STONY 
CORAL SPECIES 


NUMBER OF 
ENDEMIC 


PERCENT 
ENDEMIC 


Nihoa 


17 


4 


23.5 


Mokumanamana 


21 


8 


38.1 


French Frigate Shoals 


66 


27 


40.9 


Gardner 


29 


7 


24.1 


Maro 


41 


12 


29.3 


Laysan 


34 


11 


32.3 


Lisianski 


37 


15 


40.5 


Pearl and Hermes 


43 


14 


32.6 


Midway 


33 


8 


24.2 


Kure 


36 


15 


41.7 


Total 


80 


35 


44 



Table 4.4. Number of Rapid Ecological Assessments (REA) and Towed- 
diver Surveys (TDS) conducted by NWHI RAMP (2004 and 2006) and 
NWHI Ecosystem Reserve (2005). PHR = Pearl and Hermes. Source: 
NWHI RAMP, unpubl. data. 





2004 


2005 


2006 


REA 


TDS 


REA 


TDS 


REA 


TDS 


Mokumanamana 


3 





3 





2 


4 


French Frigate Shoals 


11 


17 


6 





10 


19 


Gardner 


3 


2 














Maro 


9 


12 


7 





9 


13 


Laysan 


3 


5 








3 


6 


Lisianski 


9 


12 








9 


12 


PHR 


14 


21 


9 





13 


26 


Midway 


9 


15 


6 





9 


15 


Kure 


9 


13 


6 





9 


13 


Total 


70 


97 


37 





64 


108 




Figure 4. 6. Percent live coral cover among reefs in the PMNM from towed- 
diver survey data, 2000-2002. Source: NWHI RAMP; map: L. Wedding 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



lar based on average estimates from 
2004 to 2006, with the highest coral 
cover at Maro (39.0%), followed by Li- 
sianski (37.5%) and French Frigate 
Shoals (26.0%; Figure 4.7). The low- 
est percent coral cover was recorded 
at Midway (1.7%), Gardner Pinnacles 
(3.5%), and Mokumanamana (4.1%). 
The lowest cover from the REA surveys 
was observed at Midway (10.1%), with 
Pearl and Hermes (12.1%) and Gardner 
Pinnacles (12.4%) also having low coral 
cover. BenthicREAswere not conducted 
at Mokumanamana from 2004 to 2006. 
Coral cover values determined from 
2002 surveys also showed the highest 
coral cover values at Maro and Lisianski 
(Friedlander et al., 2005), though their 
magnitude (>60%) was greater than the 
values derived from the line-intercept 
method in 2004-2006. 




Figure 4.7. Live coral cover among reefs in the PMNM from REA data, 
2004-2006. Source: NWHI RAMP; map: L. Wedding 



Coral cover estimates for each reef from 
towed-divers and REAs compared with 
the mean coral cover for each method 
showed higher relative towed-diver sur- 
vey values at Lisianski and French Frig- 
ate Shoals (Figure 4.8). This is likely due 
to the extensive deeper reefs that were 
not surveyed on the REAs, which are re- 
stricted to 15 m depth while towed-diver 
surveys operate down to 27 m. Higher 
relative coral cover estimates from REA 
were found at Nihoa, Midway, and Pearl 
and Hermes. The latter two locations 
have extensive back reef habitats that 
are not well surveyed by towed-divers 
due to the shallow depth. 



2.5 



2.0 



1.5 



1.0 



0.5 



0.0 



Jl 




□ REAs 

■ Tow-boards 



JH 




NEC FFS GAR MAR LAY 



LIS PHR MID KUR 



Figure 4.8. Comparisons of coral cover between towed-diver and REA esti- 
mates. Values are percent at each reef as a proportion of mean coral cover 
for that method. Source: NWHI RAMP, unpub. data. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Quantitative Estimates of Coral Genera and Species Abundance Among Reefs of the NWHI 
Relative abundance of cnidarians was assessed by computing the proportion of colonies, by taxon, that oc- 
curred within belt transects (Table 4.5, Figure 4.9). Taxa comprising more than 10% of colony abundance in 
each location are highlighted in bold type. These 2006 data surveys exemplify some general patterns seen 
in all years. The relative abundance of corals varies among regions, though Porites lobata composes a ma- 
jority of the fauna at numerous regions and is an important component of the fauna in all regions. Acropora, 
particularly A. cytherea, is an important component of the coral fauna at French Frigate Shoals, but less so in 
other regions where it occurs (Mokumanamana to Pearl and Hermes, inclusive). Pocillopora meandrina and 
Montipora capitata are both abundant in some regions but less common in others. Numerous taxa are repre- 
sented throughout the NWHI at very low levels of abundance; although 57 species of stony corals have been 
documented in the NWHI (Maragos et al., 2004), many species occur at such low frequencies that they were 
not encountered within survey transects. Thus, relatively few coral species numerically dominate throughout 
the NWHI. When species are pooled by genus, Porites, Pocillopora and Montipora collectively emerge as the 
numerically dominant genera throughout the NWHI though their relative abundance varies by region (Figure 
4.10). 



Table 4.5. Relative abundance ofcnidarian colonies in the NWHI based on RE A surveys at 64 sites conducted by NWHI 
RAMP in 2006. All cnidarian taxa for which at least one colony was tallied in at least one location are listed. Source: 
Forsman and Maragos, unpub. data. 



SPECIES 


PERCENT OF CNIDARIAN FAUNA 


MMM 


FFS 


MAR 


LAY 


LISI 


PHR 


MID 


KUR 


Acropora cytherea 


0.0% 


10.7% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


Acropora valida 


0.0% 


5.1% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


Acropora humilis 


0.0% 


0.3% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


Montipora capitata 


2.8% 


2.6% 


15.1% 


9.4% 


17.7% 


6.3% 


1.1% 


2.7% 


Montipora patula 


2.5% 


2.5% 


5.2% 


1.6% 


6.8% 


1.0% 


0.1% 


0.0% 


Montipora flabellata 


0.0% 


0.0% 


0.7% 


0.0% 


0.0% 


0.5% 


13.6% 


1.9% 


Montipora incrassata 


0.3% 


0.1% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


Pavona duerdeni 


1.9% 


2.5% 


2.5% 


3.8% 


2.1% 


0.3% 


0.0% 


0.0% 


Pavona varians 


0.1% 


0.0% 


0.0% 


2.1% 


0.1% 


0.3% 


0.4% 


0.2% 


Pavona maldivensis 


0.0% 


0.0% 


0.0% 


0.0% 


1.1% 


0.2% 


0.0% 


0.0% 


Cyphastrea ocellina 


0.0% 


7.6% 


4.8% 


4.5% 


18.8% 


1.5% 


0.9% 


1.5% 


Leptastrea purpurea 


0.6% 


1.0% 


0.2% 


0.2% 


0.3% 


7.3% 


1.1% 


3.6% 


Fungia scutaria 


0.0% 


0.0% 


0.5% 


0.0% 


0.9% 


2.4% 


0.0% 


0.0% 


Leptoseris incrustans 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


0.1% 


0.0% 


Pocillopora damicornis 


0.0% 


6.9% 


0.7% 


0.0% 


6.4% 


2.6% 


8.5% 


13.4% 


Pocillopora eydouxi 


0.0% 


0.1% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


Pocillopora ligulata 


0.7% 


0.4% 


1.2% 


0.0% 


0.6% 


0.1% 


0.2% 


0.5% 


Pocillopora meandrina 


28.9% 


8.1% 


6.5% 


17.4% 


1.1% 


26.2% 


11.3% 


52.4% 


Porites brighami 


0.6% 


1.1% 


0.3% 


3.8% 


0.6% 


0.0% 


0.0% 


0.0% 


Porites compressa 


3.8% 


15.9% 


39.8% 


1.9% 


9.7% 


8.5% 


6.3% 


5.4% 


Porites evermanni* 


2.0% 


1.5% 


1.3% 


0.2% 


11.9% 


0.0% 


0.1% 


0.1% 


Porites lobata 


55.3% 


32.2% 


20.1% 


54.9% 


20.6% 


37.1% 


55.9% 


16.3% 


Psammocora nierstraszi 


0.0% 


0.1% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


0.0% 


Psammocora stellata 


0.0% 


0.1% 


0.0% 


0.2% 


1.3% 


3.3% 


0.2% 


1.1% 


Palythoa sp. 


0.6% 


1.3% 


1.0% 


0.0% 


0.0% 


2.3% 


0.1% 


0.8% 


Total cnidarians counted 


689 


2,408 


2,443 


426 


1,920 


2,319 


1,158 


1,929 


Area surveyed (m 2 ) 


100 


500 


450 


100 


450 


650 


425 


450 


Island/Atoll abbreviations used through 

Pearl and Hermes Atoll; GAR = Gardner P 
Atoll; KUR = Kure Atoll; NIH = Nihoa Island 

* Porites evermanni is considered to be Pc 
evermanni is distinct from P. lutea. 


out this chc 

nnacles; M£ 

)/7fes lutea b 


ipter: MMM : 
R = Maro Re 

y Fenner 20C 


= Mokumanamana; FFS = 
ef; LAY = Laysan Island; L 

5, although recent molecu 


French Frigate Shoals; 
S = Lisianski Island; Ml[ 

lar analyses have confir 


and PHR = 
) = Midway 

ned that P. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Figure 4. 9. Relative abundance of coral species throughout the NWHI. Data 
are derived from colony counts within belt transects during 2006 surveys. 
Source: NWHI RAMP unpubl. data; map: L. Wedding. 



Coral Density 

Density, the number of cnidarian col- 
onies per square meter, is another 
metric of community structure that 
reflects the degree of "packing" of 
individual colonies. Such indices are 
useful when considering processes 
that may be density-dependent, for 
example, the spreading of contagious 
diseases and fertilization by spawned 
gametes during sexual reproduction. 
Colony density should be considered 
in association with size frequency dis- 
tributions in visualizing the nature of a 
population from graphed data, e.g., a 
high colony density accompanied by a 
right-skewed size frequency distribu- 
tion indicates a large number of small 
colonies, while a low colony density 
accompanied by a left-skewed dis- 
tribution indicates a small number of 
larger colonies. Density varies by tax- 
on and by habitat, as demonstrated 
for Pearl and Hermes Atoll in Figure 
4.11. 



Habitat 

Coral cover varies among reef geo- 
morphologies and reef zones (Figure 
4.12). Maro Reef has been described 
as a unique open atoll, as it lacks 
the emergent or very shallow perim- 
eter reef around a deeper lagoon that 
characterizes a classic atoll. Instead, 
the innermost area of the reef com- 
plex, with characteristics of a pro- 
tected lagoon, is separated from the 
open ocean by the surrounding mesh 
of reticulate, linear, and patch reefs. 
Average coral cover at Maro, as de- 
termined through quantitative analysis 
of benthic imagery recorded over ex- 
tensive distances surveyed by towed 
divers between 2000 and 2002, was 

14.0% (Kenyon et al., 2008a). Lisianski/Neva Shoal and French Frigate Shoals are also described as open 
atolls, the former with limited perimeter reef and the latter lacking perimeter reef to the west. Average coral 
cover at Lisianski was 19.7%, the highest average reef system value determined from extensive towed-diver 
surveys throughout the NWHI (Kenyon et al., 2007b). At French Frigate Shoals, coral cover was highest on 
the back reef (18.8%) and lowest in the lagoon (7.7%; Kenyon et al., 2006b). The most northern reef sys- 
tems in the NWHI (Pearl and Hermes, Midway, and Kure) are described as classic atolls, as their perimeter 
reefs more completely surround a central lagoon. Quantitative analysis of benthic video images recorded 
over extensive distances by towed divers between 2000 and 2003 indicates that, in these classic atoll geo- 
morphologies, coral cover is highest in the lagoon at Pearl and Hermes (19.1%) and at Kure (18.6%) and on 




Figure 4.10. Relative abundance of coral genera throughout the NWHI. 
Data are derived from colony counts within belt transects during 2006 sur- 
veys. Source: NWHI RAMP unpubl. data; map: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



the back reef at Midway (6.4%; Kenyon 
et al., 2007a; 2008b). At all three loca- 
tions, coral cover is lowest on the fore 
reef, ranging from 1.6% at Midway to 
9.7% at Kure. Coral cover is low (5.3%) 
on the shallow (<30 m) bank surround- 
ing Laysan (Kenyon et al., 2007b). 



Coral Density 
{# per square meter) 




A 



III 



h-ll 



Li 



n - 




Ll 



F* 



& 



'^-il'- ^ "^ 



Ll 



Figure 4.11. Colony density (n/m 2 ) of five coral genera at Pearl and Hermes 
Atoll, NWHI. Number of colonies (n) was determined from belt transect sur- 
veys conducted in 2002. Source: NWHI RAMP; map, L. Wedding. 



Wave exposure is a major determinant 
controlling the development of Hawaiian 
coral reefs, with moderate coral cover 
developing in areas directly exposed 
to winter wave regimes and high coral 
cover developing in sheltered embay- 
ments and areas protected from direct 
swells. In the NWHI, the greatest winter 
wave stress originates primarily from 
the northwest and secondarily from the 
northeast. At French Frigate Shoals, an 
open atoll lacking perimeter reef to the 
west, coral cover on the fore reef was 
highest in the northwest sector (approxi- 
mately 26%; Kenyon et al., 2006b). At 
Pearl and Hermes Atoll, coral cover was 
highest on the fore reef in the northwest 
and north sectors (approximately 24- 
28%; Kenyon et al., 2007a). At Midway 
Atoll, coral cover on the fore reef was 
also highest on the north sector (J. Ke- 
nyon, unpubl. data). At Kure Atoll, coral 
cover on the fore reef was highest along 
the arc extending counterclockwise from 
north to west (approximately 11-16%; 
Kenyon et al., 2008b). At Laysan, coral 
cover was highest in the northwest sector 
(7.3%; Kenyon et al., 2007b). Reef-wide 
patterns of coral cover at Maro Reef and 
Lisianski vary from those at other loca- 
tions in the NWHI with respect to wave 
exposure. At Maro Reef, an open atoll 
with no perimeter reef, the highest coral 
cover was along the northeast sector, 
closely followed by the southwest sec- 
tor (Kenyon et al., 2008a). At Lisianski/ 
Neva Shoal, an open atoll with limited 
perimeter reef, coral cover was highest 
in the southeast and southwest sectors (approximately 25-27%; Kenyon et al., 2007b). Both Lisianski/Neva 
Shoal and Maro Reef are classified as open atolls, but their complex structure of reticulate reef with little to no 
enclosure by perimeter reef likely generates a more complicated pattern of wave exposure than that experi- 
enced by atolls more clearly delineated by a perimeter reef, and accordingly they are characterized by different 
patterns of highest coral cover. 




Figure 4.12. Differences in coral cover within different reef zones in the 
NWHI. Numbers represent the average coral cover: (%) + standard error. 
Coral cover calculated from size frequency data of colonies within transects. 
Data based on REA surveys in 2002. Not all habitats were sampled at all 
reefs. Source: NWHI RAMP, unpublished data; map: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Wave Action 

High - north and west sectors experience regular large to huge wave events, especially in winter months 
where waves can exceed more than 15 m in height. Primary benthic components on exposed reefs are energy 
tolerant species and growth forms such as crustose coralline algae, non-delicate fleshy algae and encrusting 
Porites corals. 

Medium - east sectors experience nearly constant moderate wave action due to exposure from trade wind 
generated swells. Southern exposures receive occasional large swells generated by storms of tropical origin, 
especially in summer months. 

Low - the southwesterly parts of the islands generally represent the most protected sectors of the islands, 
but this generally applies to larger islands/atolls where shelter from waves is more extensive. Areas of shelter 
are minimal for the smaller islands. Atoll lagoons are the most protected of all geomorphological zones of the 
NWHI. 

Although early work in the NWHI assumed the best developed seaward reefs would occur off southwest- 
ern coasts (Grigg, 1983), analysis of imagery from more extensive and spatially comprehensive towed-diver 
surveys indicate coral cover on the fore reef is highest along the northwest, north, or northeast sectors at 
French Frigate Shoals, Maro Reef, Laysan, Pearl and Hermes Atoll, Midway Atoll, and Kure Atoll (Kenyon et 
al., 2006b; 2007a, b; 2008a, b). Only at Lisianski/Neva Shoal is coral cover highest along the southwest sector 
(Kenyon et al., 2007b) . 



Effects of Temperature on Northwestern Hawaiian Island Corals 

The coral reefs of the NWHI are in sub-tropical latitudes and exposed to large seasonal temperature fluctua- 
tions, particularly Kure, Midway and Pearl and Hermes Atolls, at the northwestern end of the archipelago. Sea 
surface temperature (SST) at these northerly atolls can fluctuate from less than 18°C in late winter (17°C in 
1997) to highs exceeding 28°C in the late summer months (29°C in 2002). Compared with most reef ecosys- 
tems around the globe, the annual fluctuations of SST of 10°C at these northerly atolls is extremely high. The 
cooler winter temperatures are thought to reduce coral growth rates. While the summer temperatures are 
generally similar along the entire NWHI, the warmest summer temperatures tend to occur at the three north- 
ernmost atolls, presumably caused by reduced mixing due to weaker winds (situated closer to the center of 
the North Pacific high pressure ridge) and decreased circulation due to large shallow water lagoons (Brainard 
et al., 2004; Hoeke et al., 2005). Roughly two-thirds of the variability in growth rate across the archipelago can 
be explained by changes in temperature and light availability (Grigg, 1982). Growth rates for a representative 
species of coral (P. lobata) dominant throughout the chain vary from 3 mm/yr to 13 mm/yr (Grigg, 1982). 

Siciliano (2005), however, reports that, while coral growth rates generally decline as a function of increasing 
latitude in the NWHI, as originally suggested by Grigg (1982), this decrease is habitat-specific. Coral colonies 
found in protected habitats throughout the NWHI chain (i.e., back reef and lagoon habitats on the atolls; em- 
bayments sheltered from wave action at the islands lacking lagoons) grow at similar rates regardless of lati- 
tude. This is may be explained by the microclimatic conditions experienced by corals growing in the shallower 
lagoon and back reef habitats, which are not closely related to offshore SST (Jokiel and Brown, 2004). Growth 
rates in these habitats may be influenced more by light and competition for space with other corals than by the 
relatively stable, and sometimes higher ambient temperatures afforded by these protected environments from 
solar heating in shallower lagoons. Conversely, corals growing in exposed habitats throughout the NWHI (i.e., 
fore reef of atolls or the reef slope of islands lacking a lagoon) experience temperatures more akin to offshore 
SST conditions, and therefore are more likely to respond to regional SST gradients, such as decreasing SST 
with increasing latitude, resulting in the measured latitudinal decrease in growth rates in these habitats. 

The growth rates reported by Grigg (1982) may also be an under-estimate at Kure Atoll because of selec- 
tive sampling. In his assessment of coral growth rate throughout the NWHI, Grigg (1982) sampled Porites 
lobata exclusively in 10 m depth from exposed southwest areas. Inspection of Porites' growth rates in three 
habitats at Kure atoll (Table 4.6, from Siciliano, 2005) reveals that the fore reef has the lowest growth rate 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 4.6. Mean linear growth rates (cm/yr) of five main reef-building coral 
genera in three habitats at Kure Atoll. Source: Siciliano, 2005. 


GENUS 


FORE REEF 


BACK REEF 


LAGOON 


Porites 


0.45 


0.74 


1.01 


Pocillopora 


1.69 


1.72 


1.69 


Montipora 


0.10 


0.10 


0.12 


Pavona 


1.63 


0.76 


0.63 


Leptastrea 


0.20 


0.22 


0.24 



for this genus, among all sites sampled. 
Therefore exclusive sampling from this 
area is likely to underestimate average 
growth rates per atoll. Even so, Pontes' 
growth rates from the fore reef habitat in 
Siciliano's study (4.5 mm/yr) are higher 
than those reported by Grigg for Po- 
rites lobata (3 mm/yr), even if the data 
for Porites exclude the faster growing, 
branching forms such as P. compressa 

(largely absent from Kure's fore reef). Encrusting and more massive growth forms of P. lobata and P. ever- 
manni, important reef builders at Kure, were included in the more recent assessment. 

Inspection of Table 4.6 also indicates that if the Porites growth rates were adjusted to reflect those of other 
reef-building corals in the fore reef habitat using Grigg's approach (i.e., by averaging Porites' growth rates with 
those of other reef-building genera), the corals' growth rate would increase to 0.8 mm/yr, rather than decrease 
as suggested by Grigg (1982), who reported an adjusted growth rate of 0.2 mm/yr for Kure Atoll. 

Monitoring Corals at Permanent Transect Sites in the Northwestern Hawaiian Islands: 2001-2006 

This section focuses on the results of 



monitoring coral communities at 27 per- 
manently marked transects established 
at seven of the 10 NWHI from 2001-2002 
and resurveyed in September 2006, and 
account for most of the 42 permanent 
transects established in the NWHI from 
2000-2002. Changes in percent coral 
cover per transect were compared be- 
tween 2001-2002 and 2006 at perma- 
nent transects (Figure 4.13). Mean coral 
cover declined by 2% from 2001-2002 
to 2006 but was not significantly dif- 
ferent between these time periods (w= 
53, p=0.51). Table 4.7 summarizes the 
results of the 2006 permanent transect 
resurveys and offers a comparisons to 
earlier surveys at the same sites. The 
changes in percent coral cover, mean 
diameter, number of coral genera and 
the density of all corals per transect are 
provided in the table. When pooling the 
data for all 27 sites, percent coral cover 
declined from 16.6 % to 14%, mean diameter declined from 22.7 to 15.5 cm, number of genera increased from 
2.9 to 4.3, and coral densities increased from 4.8 to 6.7 corals m 2 at the same set of transect sites over the four 
to five year interval. These trends are significant for all categories except percent coral cover. Changes in the 
survey techniques between the two sets of surveys can explain some of these patterns. For one, in situ census 
of corals vis-a-vis analysis of photos would likely lead to detecting greater numbers and genera for smaller cor- 
als, leading to lower mean diameter values. However, the analysis of the individual sites over the time interval 
reveals that some of these trends can only be explained by in situ observations and site-specific data. 




Figure 4.13. Percent change in coral cover between 2001-2002 and 2006 
at permanent stations in the PMNM. Source: Maragos, unpub. data; map: 
L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Table 4. 7. Percent coral cover, mean diameter, number of genera and frequency of all corals at each of 27 permanent 
transect sites surveyed in 2001-2002 and 2006. Bold numbers are the higher of two (earlier or later) values at each site. 



SITE 

NUMBER 


HABITAT 


LOCALE 


DATE 


PERCENT 
CORAL COVER 


MEAN 
DIAMETER CM 


NUMBER 
GENERA/SITE 


DENSITY 

(#/m 2 ) 


2001- 
2002 


2006 


2001- 
2002 


2006 


2001- 
2002 


2006 


2001- 
2002 


2006 


FFS- 5P 


fore reef 


NW 


7/17/2001 


19.4 


16.8 


11.6 


13.3 


4 


3 


14.2 


11.7 


FFS3P 


lagoon basalt 


N cen. 


7/16/2001 


27.7 


28.8 


19.9 


17.8 


5 


5 


8.5 


11.5 


FFS 16P 


Lagoon 


N 


9/15/2001 


4.8 


9.5 


12.3 


15.1 


5 


4 


4.6 


7.8 


FFS2P 


reef crest 


S 


7/15/2001 


27.2 


33.5 


26.2 


20.1 


7 


6 


5.3 


9.4 


FFS IIP 


back reef 


N 


10/30/2002 


39.1 


22 


76.2 


21.3 


3 


4 


1.6 


5.9 


LAY IP 


channel 


S 


9/17/2001 


5.9 


5.1 


7.7 


10.9 


3 


6 


9.5 


4.9 


LAY5P 


reef pool 


SE 


9/18/2002 


7.7 


8.7 


11 


14.6 


3 


4 


5.6 


4.8 


LIS IP 


reef crest 


S cen. 


9/30/2002 


5.3 


0.43 


14.5 


4.6 


2 


4 


3.8 


0.36 


LIS9P 


pinnacle 


E 


10/2/2002 


19.2 


24.9 


22.9 


16.3 


2 


4 


5.1 


9.8 


LIS6P 


fore reef 


N 


10/1/2002 


27.9 


46.8 


57.4 


40.6 


1 


3 


1.8 


4 


MAR4P 


back reef 


NW 


9/16/2002 


52 


34.4 


36.4 


20.6 


3 


7 


6.4 


11.4 


MAR5P 


Lagoon 


center 


9/21/2001 


4.2 


5.9 


10.9 


10.3 


2 


5 


3.8 


5.1 


MAR IP 


fore reef 


SE 


9/15/2002 


32.4 


7.9 


29.6 


9.3 


3 


7 


4.6 


8.2 


MID7P 


Lagoon 


E 


9/23/2002 


50 


1.1 


25 


7.6 


4 


4 


nd. 


3.2* 


MID 16P 


back reef 


N 


12/3/2002 


24.3 


36 


46.7 


29.7 


3 


3 


2.1 


5.8 


MID 14P 


lagoon 
pinnacle 


center 


9/24/2002 


4.7 


12.2 


8 


12 


2 


3 


12.2 


9.5 


MID 18P 


back reef 


NE 


12/4/2002 


0.7 


1.3 


6.8 


7.4 


4 


5 


2.5 


3.5 


MID 19P 


lagoon 
pinnacle 


SW 


12/5/2002 


5.1 


0.9 


14.7 


13.1 


2 


3 


2.77 


1.02 


MID 20P 


back reef 


NW 


12/6/2002 


22.3 


19.2 


48.7 


31 


3 


2 


1.46 


2.7 


MID lPa 


reef crest 


E 


12/3/2002 


3.3 


1.04 


17.6 


11.7 


2 


4 


1.73 


2.46 


MID2P 


back reef 


NE 


9/21/2002 


13.8 


13.8 


22.6 


21.3 


4 


4 


2.8 


2.8 


MID 17P 


back reef 


E 


12/4/2002 


9 


3.5 


10.4 


20 


2 


4 


4.6 


1 


MMM IP 


basalt fore 
reef 


S 


9/9/2002 


6 


14.6 


8.4 


10.6 


2 


4 


9.1 


18.4 


PHR6P 


lagoon 
pinnacle 


S 


9/19/2002 


2.53 


1.53 


11.7 


7.6 


2 


4 


1.8 


4.8 


PHR7P 


lagoon patch 
reef 


center 


9/27/2002 


24 


20.7 


25.6 


10.1 


1 


3 


4.64 


19.8 


PHR9P 


Pass 


S 


9/28/2002 


1.69 


0.23 


14.4 


9.5 


2 


4 


1.07 


0.4 


PHR12P 


fore reef 


SW 


9/29/2002 


8.95 


7.11 


15.5 


11.1 


3 


6 


3.15 


6.48 


TOTALS 


27 




MEANS 


16.64 


14 


22.69 


15.46 


2.9 


4.3 


4.8 


6.7 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Mokumanamana 

Permanently marked transect NEC-IP 
is located in a semi-sheltered recessed 
bay or bight off the southwest side of 
Mokumanamana and is the only perma- 
nent transect established to date. The 
site was re-surveyed on September 3, 
2006. All coral population parameters in- 
creased during the four year period be- 
tween the two surveys. Mean diameter 
increased from 8.4 cm to 10.6 cm. Coral 
densities increased from 9.1 to 18.4/m 2 , 
generic diversity increased from two to 
five genera, and coral cover more than 
doubled to 14.6%. All four of the smaller 
size classes for total corals increased 
substantially during the four year period 
although there were only a few corals 
represented at the higher size classes 
for each survey period (Figures 4.14 and 
4.15). Overall, all smaller size classes in- 
creased and the larger stayed the same. 
Severe exposure to waves from any di- 
rection and the large winter swell from 
the northwest may prevent development 
of large, high profile corals. The lobe 
coral (Porites lobata) followed by the 
rose/cauliflower coral (Pocillopora me- 
andrina) continue to dominate the coral 
fauna, although the zoanthid soft coral 
(Palythoa tuberculosa) has emerged as 
a common coral on the transect in 2006, 
even though absent in 2002. The num- 
ber of Porites corals doubled for the all 
four smaller size classes while Pocil- 
lopora also increased dramatically in all 
size classes. Despite the exposed and 
scoured environment at the site, corals 
have increased dramatically at NEC-IP 
over the four year period. 



Coral Percent Change 


■ 


-Q.98& 


- -0.6S% 


O 


-rj.67% 


-4.34% 


© 


■G.33% 


-0.13% 


• 


0.14% - 


O.flffft 


• 


0.&7% 


IJMI 



Banthic Habitat Type ~J Ham&oflorn, indeterminate c 

| Hard&otlom. >1 0% oruslose coralline algae ^^H hartfbctLom uncotanized 
^| Hard bottom, >1 0% live coral ~~\ Land 

| HardbDHom, >1 D% macrnslgae ^| Unconsolidated 



| Unconsolidated with * MWS macrQatgaa- 
Unclassified N 



A 



164* WW 



iet-42-vi 



Figure 4.14. Percent change in coral cover at Mokumanamana between 
2002 and 2006. Source: Maragos, unpub. data; map: L. Wedding. 




v 3 ^ ^ / / 

to V" 1> t>> 



•v* <? / / -f 

10 V" T> t>> 



Coral Size Distribution (cm) 



Figure 4.15. Changes in the proportion of each genus, size distribution and 
cover for corals reported at Mokumanamana Island permanent sites be- 
tween 2001-2002 and 2006 permanent site IP. Source: Maragos, unpub. 
data. 



French Frigate Shoals 

Eleven permanent monitoring transects for corals were established at French Frigate Shoals from 2001-2002, 
with four resurveyed in September 2006. These are: Serendipity Hollow (site FFS IIP) on the northern back 
reef; site FFS 16P at the CREWS buoy site in the northern lagoon; site FFS 3P off the north side of La Perouse 
Pinnacle in the north central lagoon; and site FFS 2P just west of Disappearing Island off the south reef crest 
of the atoll. Large northerly swell and limited time during the visit prevented safe access to and survey of the 
remaining seven sites. 

All sites showed increases in coral density (number of corals/m 2 ) and most showed increases in coral cover 
and generic diversity (Figures 4.16 and 4.17). The northern lagoon site (FFS-16P) showed increases in mean 
coral diameter and dramatic increases in the other coral parameters between 2001 and 2006, although the 
three smallest size classes peaked in 2002. The Serendipity Hollow (FFS IIP) site appears to have ex- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



perienced wave damage in the recent 
past, based upon broken and over- 
turned corals observed at the site in 
2006, but coral populations continued 
to maintain high values and post mod- 
est increases in all size classes except 
the largest two. The La Perouse site 
(FFS 3P) posted increases in generic 
richness, with the addition of Acropora, 
Montipora and Psammocora, although 
there were slight decreases in the abun- 
dance of Pocillopora and Porites. Cor- 
als at the Disappearing Island, site (FFS 
2P) showed consistent gains in the four 
smallest size classes and dramatic in- 
creases in the number of genera over 
the five year period. Moreover, six coral 
genera noticeably increased in abun- 
dance (Acropora, Pocillopora, Porites, 
Palythoa, Pavona, Psammocora) with 
Montipora showing a modest decline. 



Coral Percent Changs 


• 


-e.ee* 


• -e.ee* 


• 


-0.67* 


-0 34* 


• 


-0.33* 


-0.13% 


• 


0.14*. 


o.ee* 


• 


0.87*. 


ua 



Bonthic Habitat Typ* 

B HardboLlom. >1C% crtwlose coralline a 
| Katfbottom, >10% he coral 
^ HardtwQem, >10% iri9CfQ?i9S9 




| UiKonBoidBlBd 



166 D 24'W 



Figure 4.16. Percent change in corai cover at French Frigate Shoais be- 
tween 2002 and 2006. Source: Maragos unpub. data; map: L. Wedding. 



n Acropora 
■ Porites 



■ Leptastrea □ Montipora □ Pocillopora 

□ Palythoa D Pavona □ Psammocora 



40 
30 
20 
10 



^A 








I 






□ Cyphastrea 
D Pavona 




J r>° ^ .>° ,»° V* & 



n Montipora 
■ Porites 



£ 



1 






^ <$ $> 



Coral Size Distribution (cm) 



□ Acropora 

□ Pavona 



D Cyphastrea D Montipora D Palythoa 
□ Pocillopora ■ Porites □ Psammocora 




u 



■ o- t> & ^ 



ll 





D □ Acropora D Montipora 


□ Pocillopora ■ Pontes 


140 - 






120 - 


2001 - 2002 




2006 


o o 

O CO 

(ooo't;) 








E 








V 60- 








C6 
0) 








< 40 - 




1 




20 - 
- 


m m 


1 


--■III 



jr ,-v K )» K # ^ & 



*> •> T> t> oty N 



Coral Size Distribution (cm) 



Figure 4.17. Changes in the proportion of each genus, size distribution and cover for corals reported at French Frigate 
Shoals permanent sites between 2001-2002 and 2006: a) 2P, b) 3P, c) 16P and d) IIP. Source: Maragos, unpub. data. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Maro Reef 

Three permanent transect sites were 
established at Maro Reef between 
September 2001 and September 2002 
(Figures 4.18 and 4.19). Site MAR IP is 
located on a southeast facing fore reef 
of a patch reef. Site MAR 4P is located 
at the far northwest end of Maro, on a 
south-facing leeward fore reef. MAR 
15P is near the middle of these two ex- 
tremes in the central shallow protected 
lagoon 20 m north of the NOAA CREWS 
buoy. All sites were resurveyed on 7-9 
September 2006. Wave exposure varies 
considerably among the three, with the 
southeast site most exposed, the cen- 
tral site least exposed, and the north- 
west subjected to intermediate wave but 
stronger current exposure. 



Coral Percent Change 


• 


-0.9B* 


-■MM 


• 


-0.67% 


--0.34% 


• 


-0.33% 


-0,13% 


• 


0.14* 


0.85* 


• 


0,87% 


1.8* 



lit-} 






•j£ 






rv 







Bcnthic Habitat Type 
| Hardbmlom, >10%t;ruskisy rary'lmea 
| HirdbonDm h >10%1hr«cortf 
HardboflBm. >1Q% macrcfllgae 



Ha-dbollom 



_] UrKonsol-dated with < %0%- mBcroalgao 



Um.i 
I Unconsolidated 



Ai 



170'42'W 



Figure 4.18. Percent change in coral cover at Maro Reef between 2002 and 
2006. Source: Maragos, unpub. data; map: L. Wedding. 



Coral populations showed mixed trends 

over the four to five year period. The 

number of genera increased from 2-3 per site in 2001-2002 and to 6-7 per site in 2006. The dominant corals 

at all three transects were the lobe coral (Porites lobata) and the rose coral (Pocillopora meandrina). Other 

common coral genera were Pavona and Montipora, the latter absent at the southeast site in 2001 (MAR IP), 

but more abundant at all three in 2006. The encrusting brain coral (Leptastrea pruinosa) was absent at all three 

sites in 2001-2002 but emerged as a common species at the northeast site (MAR 4P). 



nPsammocora 



■ Porites O Pocillopora OMontipora DCyphastrea 



I 






LL 



n 



B 

70 

60 

& 50 

o 

o 

d 40 

30 

CO 

1 20 

10 




UZoanthus UPorites nPocillopora nPavona UPalythoa UMontipora UCyphastrea 
2001- 2002 2006 

■ MM -■■! 



J* ^ k> Q V* N*° 



\"> f 









^ J* K& ^ ^ ^ ^ 

* N N <P ^ & 1 






Coral Size Distribution (cm) 



Coral Size Distribution (cm) 



400 

350 

300 

§ 250 

■^ 200 
u 

B 150 
0) 

* 100 

50 



UPorites nPocillopora nPavona UPalythoa UMontipora ULeptastrea nAcropora 



2001 - 2002 



^JSl 



ll 



-,-,H,B,», 



S P >$> * ^ # 5? 
« O' T>' ^ #" 



Coral Size Distribution (cm) 



Figure 4.19. Changes in the proportion of each genus, size distribution and cover for corals reported at Maro Reef perma- 
nent sites between 2001-2002 and 2006: A) 15P, B) IP and C) 4P. Source: Maragos, unpub. data. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Mean diameters decreased at all sites, from 10.9, 29.6 and 36.5 cm in 2001-2002 to 10.3, 9.3 and 20.6 in 2006 
at sites MAR IP, MAR 4P and MAR 15P, respectively. However, coral densities (number of corals/m 2 ) increased 
at all sites over the same period from 3.8, 4.7 and 6.4 to 5.1, 8.4 and 11.2 corals in 2006 at the same respective 
sites. Coral cover decreased at MAR 4P and IP and increased at MAR 15P At site MAR 15P the numbers of 
smaller corals generally increased, but at the southeast site (MAR-IP) and the northwest site (MAR 4P) the 
numbers for all size classes decreased. Many larger corals appeared to show signs of fragmentation to smaller 
sized corals at the two more exposed sites (MAR-IP, MAR-4P) that could explain the observed shifts in size. 



Laysan Island 

Two permanent transect sites were es- 
tablished at Laysan Island in Septem- 
ber 2002 and resurveyed in September 
2006. Site LAY-IP is located along the 
north side of the sand channel in the 
southwest embayment (Figures 4.20 
and 4.21). Site LAY-5P is located on 
the shallow southwest reef crest, just 
south of the embayment and west of the 
NOAA monitoring buoy. Both transects 
are sheltered from heavy winter swells 
from the northwest Pacific that strike the 
NWHI during the winter months. 



Coral Percent Change 


• 


-0.96% 


i)«fi'.<- 


• 


■o.g™ 


■ -0.34% 


• 


•0,35% 


•013* 


• 


0.14K • 


0.96% 


o 


Q.B7%- 


1jfi% 



Site 




■W 



- 



Benth i c Habitat Type 

'10% cnjslo&B cora 
'10% Ibvft OOfSl 
^ Hardbcflom. >10% mserwilgae 



| HgrdtwUom. mcJele*mingte ppygf 
- ^^| Hardboitom uncolomzfid 
1 Land 

3 



^2 UncoratmdMM win « 10% macroaijM 
unclassified N 

0.S 1 2.._ A 



Coral population parameters showed 

mixed trends over the four-year period. 

Generic richness increased from three 

to four genera at sites LAY-IP and LAY- 

5P from 2002-2006. Mean diameters 

also increased at both sites, from 7.7 

cm to 10.9 cm at LAY-IP and from 11 

cm to 14.7 cm at LAY-5. However, coral 

densities dropped at both sites, dramatically at LAY-IP from 9.5 to 4.9, and less so at LAY-5P from 5.6 to 4.8 

over the four- ear period. About 10% of the corals, mostly Porites lobata, at LAY-1P appeared sick or dying, 

and overall coral health at this site appears less than reported there in 2002, including a 50% decrease in coral 

cover. There were few sick or dying corals at the reef crest site LAY-5P although coral cover declined slightly. 



Figure 4.20. Percent change in coral cover at Laysan Island between 2002 
and 2006. Source: Maragos, unpub. data; map: L. Wedding. 



A 


B 

DCyphastrea MLeptastrea DMontipora DPavona DPocillopora ■Pontes 




DCyphastrea OPavona OMonlipora nppcillopora ■ Pontes 


14 - 






18 ■ 




12 - 




16 - 








„ 




_ 


2001 - 2002 


2006 




§ 10 - 


2001- 2002 2006 


o 14 - 
o 








*H 




=■ 12 - 








| 8 - 




E 

" 10 - 


■ 


■ 


m 


Area 


i ■ ■ l l 


n 

a) 

< 8 - 








4 - 


I I 


6 - 
4 - 








- 


.Mill ill 


2 - 


.■II 


. 1 
1 ill 




* ** <? / *?v / ^ «* ** / // ^ 




Coral SizeDistribution (cm) Cora , size Distribution (cm) 


Figure 


>s 4.21. Changes in the proportion of each c 


enus, size distril 


wtion, and cover 


for corals reported at Lay. 


;an Island 



permanent sites between 2001-2002 and 2006: A) IP B) 5P. Source: Maragos, unpub. data. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Laysan's smaller oval shape renders it 
vulnerable to severe exposure to waves 
from the west to northeast, and the large 
winter swell from the northwest may 
prevent development of large, high pro- 
file corals at shallow depths. Although 
not measured during the 2006 surveys, 
benthic algal cover appeared especially 
high at the west channel site LAY-1P, 
and algae is prolific at other sites (J. 
Maragos, per. obs.). Aside from its small 
size, Laysan Island supports over a mil- 
lion breeding seabirds and significant 
monk seal and sea turtle populations. 
Together with seepage from its high 
nutrient-laden hypersaline lake, Laysan 
Island may be subsidizing substantial 
localized productivity from local nutri- 
ents which in turn may favor the growth 
of algae over corals in most shallow reef 
habitats. 



Coral Percent Chang* 


• 


-0.98% 


--&68S 


• 


«an 


■-0.34% 


• 


-0.33% 


0.13% 


• 


0.H* ■ 


0WV. 


• 


0.07% - 


16% 






















Benthic Habitat Type 

■ Hardbonom. >10% cnrttose coralline 
B HordooElom, >10% irvo coral 
H HardtwHcwn, >10% ma«703Jg9e 


i(gH 


I Hardbattcm, Indeterminate 
^^| HardboMom. uneoimiized 
■ Land 

| UnconsDlkjacsd 


cover 


1 UnawiKiadeled win 
UndassffiBd 
3.5 S 


= 10% rnwnaelgae 
N 



Figure 4.22. Percent change in coral cover at Lisianaski-Neva Shoals be- 
tween 2002 and 2006. Source: Maragos, unpub. data; map: L. Wedding. 



Lisianski Island 

Three permanent 50 m long transects 
were established in shallow water near 
Lisianski Island in September-October 
2002 and all three were resurveyed 
in September 2006 (Figures 4.22 and 
4.23). Site LIS 1P was established 
on the shallow southwest fore reef at 
depths of 1 .2 to 4.6 m. Site LIS 6P was 
established near the opposite, east side 
of the island at similar depths along the 
fore reef-reef crest margin. The third site 
LIS 9P was established off the northern 
fore reef slope of the island at depths of 
7.3-14.9 m. Water visibility, wave action 
and currents are notably stronger at the 
latter two sites (LIS 6P and 9P) located 
windward of the island. Site LIS 1P is 
down-drift of the island and in the direct 
path of current and water flow from the 
island. 

In 2002 coral coverage was estimated 
at 5.3%, coral density at 3.8 corals/ 
m 2 , and mean diameter at 14.5 cm at 
the southwest site LIS-1P Corals there 
were primarily plates of Montipora cor- 
als intermixed with green algal growths 
of Neomeris and Microdictyon. On the 
opposite, eastern site, LIS-6P, large 
plates of the same coral Montipora cf. 



B 


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o 


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□ Pocillopora 


■ Pontes 






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8 - 


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6 - 














5 - 
















4 - 
















3 - 




















2 - 
1 - 
- 

























n° ^ tP <j? .# 

fc *' n> * „>- 



DCyphastrea D Montipora D Pocillopora ■ Pontes 



2002 



2006 



- . n 



o 



i i 



^ */ ^ >? f s« 






DCyphastrea D Montipora D Pocillopora 



2002 



n 



2006 



n 



n 



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W K K' K' a> 



v * ^ J? J? & J? 



Coral Size Distribution (cm) 



Figure 4.23. Changes in the proportion of each genus, size distribution, and 
cover for corals reported at Lisianski-Neva Shoals between 2002 and 2006: 
A) IP, B) 9P and C) 6P. Source: Maragos, unpub. data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

turgescens dominated the bottom, averaging 27.9% coral cover, 1.8 corals/m 2 , and with an average diameter 
of 57 cm in 2002. The north, offshore site consisted of hard bottom covered with encrusting Montipora corals 
at shallow (6-7.6 m) depths and transitioned to larger, widely spaced colonies of Porites lobe corals and Pocil- 
lopora rose corals over a rubble sand bottom at depths of 9.1-15.2 m. Coral coverage was 19.2% with a mean 
diameter of 23 cm and frequency 5.06 corals/m 2 in 2002. 

Four years later, resurveys yielded a near total collapse of corals at site LIS-1P. Coral cover was estimated at 
0.43%, mean diameter at 4.56 cm, and coral density at 0.36 corals/m 2 , translating to only 17 corals in the three 
smallest size classes for the entire transect. In great contrast, the two windward sites fared much better over 
the four years with coral cover higher at both in 2006. Mean diameter at site LIS-6P was less but still substan- 
tial at 40.6 cm. Likewise site LIS-9P showed smaller mean diameter for corals at 16.3 cm, but coral frequencies 
nearly doubled at both windward sites, with 2006 values at 4.00 and 9.84 corals/m 2 respectively. The small 
brain coral Cyphastrea appeared at all three sites for the first time in 2006, and Pocillopora increased in size 
and numbers at all three sites. The plate coral Montipora nearly disappeared by 2006 at site LIS-1P with only 
10 small colonies remaining from 184 mostly larger colonies in 2002. At the other two sites Montipora plate 
coral increased dramatically for smaller to middle sized corals. The lobe coral Porites also declined at the 
southwest site LIS-1P, but increased modestly at the northern site LIS-9P Figure 24 illustrates the changes in 
size distribution for all corals over the four year period at the three sites. 

The collapse of the coral community at southwest site LIS-1P may have been caused by a coral bleaching 
event followed by the overgrowth of algae, the latter perhaps stimulated by plentiful nutrients leaching from 
the island and derived by extensive guano production generated by the large resident seabird populations at 
Lisianski. Ambient water temperature was higher at the site during the 2006 survey and may have a longer 
residence time in shallow depths where solar heating would be higher. Also, the waters to the southwest were 
noticeably greener due to higher phytoplankton productivity. In contrast, water clarity was better, water motion 
stronger and temperatures cooler at the two windward sites that showed increased coral development. 



Pearl and Hermes Atoll 
Eight permanent coral transects were 
established at Pearl and Hermes prior to 
the September 2006 visit. Four of these 
were established at the site of the 2000 
grounding of the fishing vessel Sword- 
man I. The 2006 REA team did survey 
one or more of the back reef sites but 
these data are currently not available. 
The remaining four permanent transects 
established in 2002 consist of site PHR 
6P, a shallow south lagoon pinnacle 
slope; PHR 7P, a central lagoon patch 
reef in the finger coral gardens; PHR 
9P on the floor of the main south pass 
of the atoll; and PHR-12P off the south- 
west fore reef of the atoll (Figures 4.24 
and 4.25). 

Coral populations showed smaller mean 
diameters (7.6 to 11.1 cm) in 2006 com- 
pared to their corresponding 2002 values 
(11.7 to 25.7 cm) at all sites. However, 
all sites showed higher generic diversity 
ing 2002 values (one to three genera), 
except PHR-9P, varying from 0.4 tol9. 



Coral Percent Changs 


• 


-Q.SB% 


- -0.66% 


O 


■0.67% 


--0.34% 


• 


■0.33% 


-0.13% 


• 


0.14% - 


o.as% 


• 


07% - 


16% 




Br:rHln<; Habitat Type ^J HardbollDm. indeterminate «*Bf 
1 H3«Jbeflwn. >tO%dvsl&secwanineflagae ^^ hardbotlorn, unedamwd 


J U neon sol Ida ted with < 

UnCass'ined 


10% macros Igae 
N 


^| Hardbqltom,. >10% live canal 1 Land 

| HBrdtxHtom, >10%macrDalgBB Unconsolidaled 


2 4 


A 







Figure 4.24. Percent change in coral cover at Pearl and Hermes Atoll be- 
tween 2002 and 2006. Source: Maragos, unpub. data; map: L. Wedding. 

levels in 2006 (three to six genera/transect) compared to correspond- 
The 2006 coral populations also showed larger densities at all sites 
8 corals/m 2 , compared to 2002 levels (1.07 to 4.64 corals/m 2 ). The 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

small brain coral (Cyphastrea ocellina) was reported on three transects for the first time in 2006, and Palythoa, 
Pavona and Psammocora were all reported for the first time on one transect each. 

The three smallest size classes dominated the coral numbers at all 2006 sites except the south pass site 
(PHR-9P). Coral populations in the pass were much diminished from their 2002 levels at all size classes. 
During both the 2002 and 2006 surveys, currents ranged from 2 to 3 knots in the pass, with considerable sus- 
pended sediments. Scour and periodic wave action may be controlling coral development in the south pass. 
At the fingercoral garden site (PHR-7P), corals in the three smallest size classes were an order of magnitude 
more abundant in 2006 compared to 2002 levels for Porites compressa, although there was no substantial dif- 
ference in the larger four size classes. In general, all corals combined were more numerous at the larger size 
classes at all four sites in 2002, Figure 4.25 compares size distributions for all corals in 2002 and 2006 at the 
four permanent coral transect sites. 



7 
6 

s- 5 

o 
o 

~E 
? 3 
(B 

0) 

< 2 

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ILeptastrea DPocillopora ■ Porites D Psammocora 



ILeptastrea nPocillopora ■ Porites D Psammocora 



: 



r 



J 



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~ 6 

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td 3 

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5 2 

1 




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xa 



<b' 






I&- 



V 



e 



,"*■ 



..V 



& 



Leptastrea UMontlpora □ Pavona nPocillopora 



» 



J 1 ^ 



□ Cyphastrea □ Pocillopora 



7 

§6 

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H 

5» 5 

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CTJ 

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< 

2 

1 




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^1 



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1 



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fo -> «1> k> c s> 



$> $> $> <£> 

<yV T>' t^" 



W M n > ^ J* -7* 



% v f 



V 6 „>° J? JP .-* # v* 



Coral Size Distribution (cm) 



Coral Size Distribution (cm) 



Figure 4.25. Changes in the proportion of each genus, size distribution, and cover for corals reported at Pearl and Hermes 
permanent sites 6P, 9P, 12P and 7P from 2002 and 2006. Source: Maragos, unpub. data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Midway Atoll 

Eleven permanently marked transects 
were established at Midway Atoll Na- 
tional Wildlife Refuge between 2000- 
2002. Eight sites were resurveyed in 
September 2006 and are shown in Fig- 
ures 4.26 and 4.27. Five of the resur- 
veyed sites are situated on the northern 
and eastern back reefs (MID lPa, 2P, 
7P and 17P), two on the NW back reef 
(MID 18P, 20P), one near the southwest 
reef crest (MID 19P) and one on a cen- 
tral lagoon patch reef (MID 14P). 



Coral Percent Change 




■ -D.9B% - -GJEffft 




• HIST* ■ -CM% 




# -fias% -om 


V ' -^ 


Q.14%-0.tt% 




^ 0.67'/, -1.6% 




j!** JK.*J 




Benlhic Habitat Typa 



• 1 0% crustose coral 
| HawlboltorTi. >1Q% IN* coral 
j Hanjbcflorn. >10% macfMlg» 



1 htardboaom. Indeterminate cover T i 

* H; hiardbchom, uncolnruzed [ 

^] Land 1 



A 



Figure 4.26. Percent change in coral cover at Midway Atoll between 2002 
and 2006. Source: Maragos, unpub. data; map: L. Wedding. 



n Pocillopora ■ Pontes O Psammocora 



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*£ 4 

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(B 3 
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□ 



1 




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Coral Size Distribution (cm) 

□ Pocillopora ■ Pontes D Psammocora 



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Coral Size Distribution (cm) 



DCyphastrea DMontipora D Pocillopora MPorites 



Li 



ul 



J 3 ,n? JP j£ 

<° »> 1> t> 



2006 



2006 



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Coral Size Distribution (cm) 



Coral Size Distribution (cm) 



Figure 4.27. Changes in the proportion of each genus, size distribution, and cover for corais reported at 7 permanent sites 
at Midway: A) MID lPa, B) 2P, C) 14P, D) 17P, E) 18P, F) 19P and G) 20P Source: Maragos, unpub. data. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



E 
2.5 



sf 1.5 



70 
60 
50 
40 
30 
20 
10 




HCyphastrea n Montipora MPalythoa HPavona n Pocillopora ■ Pontes 






1? IP 

Coral Size Distribution (cm) 



V «= n> o> i> 



0- v t>' 



D Montipora D Pocillopora ■ Pontes 



H 



n 



R 




Coral Size Distribution (cm) 



« V A" nV" A" ^ -^ 



v ^ v t* v o,v 



V 6 o>° ^ .>° ^ v* v fo0 



V ^ v t> v p>' 



Coral Size Distribution (cm) 



Figure 4.27 (continued). Changes in the proportion of each genus, size distribution, and cover for corals reported at 7 
permanent sites at Midway: E) 18P, F) 19P and G) 20P. Source: Maragos, unpub. data. 




Figure 4.28. In 2006 both Cyphastrea ocellina (left) and Psammocora stel- 
lata (right) were reported for the first time at Midway Atoll. Photos: J. Mara- 
gos. 



Overall, coral populations showed mixed 
trends during the four year period. The 
abundance and numbers of Porites 
lobata and P. compressa decreased 
slightly. The rose coral Pocillopora spp 
showed no clear trend, and the abun- 
dance of Montipora capitata and M. cf. 
turgescens generally increased from 
2002-2006. The small brain coral (Cy- 
phastrea ocellina) and small branch- 
ing coral (Psammocora stellata) were 
reported for the first time at several 
transects in 2006 (Figure 4.28). Overall there were no trends for coral mean diameter or frequencies over the 
four year period, although generic diversity per transect increased at most sites. 

The eastern back reef site MID-lPa showed increases at smaller size classes but declines in larger corals. 
The neighboring east back reef site 17P showed major declines in all size classes and genera. Finger coral 
(Porites compressa) at the southeast lagoon site MID-7P were abundant in 2002, but in 2006 were all dead 
but still standing. The southwest reef crest/back reef site MID-19P also showed catastrophic declines in all 
corals. In contrast, the two northern back reef sites MID-20P and MID-18P dominated by Montipora showed 
major increases over the four year period. In 2002, Montipora at these sites were especially hit hard by coral 
bleaching, but now have rebounded to levels comparable if not higher than those reported earlier. The reasons 
for the major coral declines at sites MID-7P, -IP and -17P are not clear but warrant continued monitoring. The 
declines may be related to the residual effects of WWII and post WWII military construction at Midway, which 
resulted in the dredging of a deep channel between the lagoon and ocean and the likely change in lagoon 
circulation and water levels. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Descriptive Statistics for Corals from 2006 Rapid Ecological Assessment Data 

Mokumanamana 

Table 4.8. Descriptive statistics for corals from 2006 REA data at Moku- 



In 2006 only two stations were sampled 
at Mokumanamana Island (Table 4.8, 
Figure 4.29). Average coral species 
richness (x = 11.5, SD ± 2.1) ranked 
the highest among all reefs surveyed. 
Percent live coral cover was 22.1% (SD 
± 15.9) and ranked forth overall. Den- 
sity of coral colonies (number/m 2 ) also 
ranked first among all locations sampled 
in 2006 with a mean density of 6.9 colonies nr 



manamana Island. Source: NWHI RAMP, unpub. data. 



CHARACTERISTIC 


NUMBER 


MEAN 


STD 
DEV 


STD 
ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Richness 


2 


11.5 


2.12 


1.5 


-7.56 


30.56 


Coral cover (%) 


2 


22.06 


15.94 


11.28 


-121.2 


165.32 


Density (no. m 2 ) 


2 


6.89 


0.55 


0.39 


1.93 


11.85 



ttreww 




Coral Sp«cles Richness 


■ 


2-4 




• 


5-7 




O 


8-9 




• 


10-11 




• 


12-14 
: .11- i 


N 

A 





02 0* 



lerasrw 



Percent Corel Cover 
■ to-55 

• 5 6 - 1Z0 

• 12.1-26.0 

• H.I -38.2 

• 363-696 

WdLw *2fl m f\ 
0.2 <M 


^TKr~w~- 









i&r«'ww 




164N2 , M"W 



1W4V24-W 



Figure 4.29. Mokumanamana coral statistics and distribution for 2006. Species richness (top left), percent coral cover (top 
right) and coral density (number m 2 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



French Frigate Shoals 
French Frigate Shoals ranked third over- 
all in coral species richness with an av- 
erage of 9.9 coral species per transect 
(Table 4.9, Figure 4.30). Percent live 
coral cover was 29.7% and also ranked 
third among all locations surveyed in 
2006. Coral colony density was 4.82 
m 2 and again ranked third among loca- 
tions. 



Table 4.9. Descriptive statistics for corals from 2006 RE A data at French 
Frigate Shoals. Source: NWHI RAMP, unpub. data. 



CHARACTERISTIC 


NUMBER 


MEAN 


STD 
DEV 


STD 
ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Richness 


10 


9.9 


3.07 


0.97 


7.7 


12.1 


Coral cover (%) 


10 


29.71 


18.68 


5.91 


16.3 


43.07 


Density (no. nr 2 ) 


10 


4.82 


2.37 


0.75 


3.12 


6.52 



• • 




• 




Coral Spoclfts 


Richness 


- 


2-4 




• 


ft I 




• 


8-9 




• 


10-11 




• 


12-14 




1 


Urnl 
Waters 


. A 












lfYTW)l()rK 




Perctni Com Covtr 


- 


1.0- 


5.5 







5.6- 


12 C 




O 


-.2 1 


-26.0 




O 


2B.1 


-38.2 




O 


36 3 ■ 69,6 

Land 
V-.i— -.-: ■"■ 


N 

A 


1 


2 


4 







Figure 4.30. French Frigate Shoals coral statistics and distribution for 2006. Species richness (top left), percent coral 
cover (top right) and coral density (number m 2 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Maro Reef 

Species richness at Maro Reef was 9.1 Table 4.10. Descriptive statistics for corals from 2006 REA data at Maro 
per transect and ranked fourth overall Reef - Source: NWHI RAMP ' un P ub - data - 
(Table 4.10, Figure 4.31). Coral cover 
was high (x = 33.3, SD ± 17.8) and 
ranked second overall. Density was the 
second highest among all sites and av- 
eraged 5.43 colonies/nr 2 . 



CHARACTERISTIC 


NUMBER 


MEAN 


STD 
DEV 


STD 
ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Richness 


9 


9.11 


1.45 


0.48 


7.99 


10.23 


Coral cover (%) 


9 


33.29 


17.85 


5.95 


19.6 


47.01 


Density (no. m 2 ) 


9 


5.43 


1.47 


0.49 


4.3 


6.56 





• 

9 


Cofsl Speeds Richness 

■ 2-4 

■ 5-7 

• 8-9 

• 10 -If 
9 12-H 

1 
VftlDr <20 m ^ 

DIE 4 


• 

• 
• 







Percent Coral Cover 


■ 


1.0-5.5 


• 


5.S - 12.0 


• 


12.1-23.0 


• 


26.1 -33 2 


• 


38 .3 ■ 69 6 

Uuid N 
NVnlDT 120 m f\ 


L 


1 


2 4 







Coral Density 




• 06-20 




• 2.1-3.? 




$ a a -46 




• 4,7 -sa 




# R»-toa 




. and 


N 

A 


D 1 2 <\ 




Figure 4.31. Maro reef coral statistics and distribution for 2006. Species richness (top left), percent coral cover (top right) 
and coral density (number m 2 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Laysan Island 

Lasyan Island had limited sampling ef- 
fort in 2006 (Table 4.11; Figure 4.32). 
There were an average of 8.0 species 
recorded from transects, ranking fifth 
overall. Coral cover was also intermedi- 
ate (fifth in rank) with an average cover 
of 17.7%. The density of coral colonies 
was low (seventh in overall rank) with 
an average of 2.8 coral colonies/nr 2 . 



Table 4.11. Descriptive statistics for corals from 2006 REA data at Laysan 
Island. Source: NWHI RAMP, unpub. data. 



CHARACTERISTIC 


NUMBER 


MEAN 


STD 
DEV 


STD 
ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Richness 


3 


8 


1 


0.58 


5.52 


10.48 


Coral cover (%) 


3 


17.65 


12.86 


7.42 


-14.3 


49.59 


Density (no. m 2 ) 


3 


2.84 


1.06 


0.61 


0.2 


5.48 



i 

z 

I 




• 
• 






Coral Speciss Richness 

• 2-4 

• 5-7 

• B-9 

• 10-11 

• 12-14 

W*Sf<ZDrrt f\ 
■•■ •■> 5 1 2 

















• 


z 

1 

■ 


Percent Coral Cower 
- 1.0-5.5 

* 50-12,0 

• 12.T-26.Q 

# 2SJ-3B.2 

# 39 2 - £9.0 

I 

o os i ;., 










Figure 4.32. Laysan Island coral statistics and distribution for 2006. Species richness (top left), percent coral cover (top 
right) and coral density (number m 2 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Lisianski. Source: NWHI RAMP, unpub. data. 



Lisianski-Neva Shoals 

Lisianski-Neva Shoals averaged 10.8 Table 4.12. Descriptive statistics for corals from 2006 RE A data at 
coral species per transect and ranked 
second overall (Table 4.12, Figure 4.33). 
It ranked first in coral cover with an aver- 
age of 36.7%. Density of coral colonies 
was intermediate with 4.3 colonies/m 2 . 



CHARACTERISTIC 


NUMBER 


MEAN 


STD 
DEV 


STD 
ERR 
MEAN 


LOWER 

95% 


UPPER 

95% 


Richness 


9 


10.78 


2.28 


0.76 


9.03 


12.53 


Coral cover (%) 


9 


36.74 


20.77 


6.93 


20.8 


52.71 


Density (no. m 2 ) 


9 


4.27 


1.54 


0.51 


3.08 


5.45 







• 




■r Q J^t 




m 




• 




cwal Species Richness 


• 
• 


■ 2.4 
> 6-7 

• a-s 
#1 10-11 

# 12-14 


* • 


- !l ~ A 

12 -L 















Ink-* 9 

o 

O 




Coral Density 

• 0.6 - Z.D 
■ 2.1 - 3.7 

• 3.B - 4.G 
% 4.7-5.B 
5.9-10.2 

12 4 









Figure 4.33. Lisianski-Neva Shoals coral statistics and distribution for 2006. Species richness (top left), percent coral 
cover (top right) and coral density (number m 2 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Midway Atoll 

Species richness of corals at Midway 
was the lowest of any reef surveyed in 
2006 with an average of four species 
per transect (Table 4.13, Figure 4.34). 
Coral cover was 13.8% and ranked sixth 
overall. The density of coral colonies 
was 2.7/nr 2 , which was also the lowest 
among all location. 



Table 4.13. Descriptive statistics for corals from 2006 REA data at Lisian- 
ski. Source: NWHI RAMP, unpub. data. 



CHARACTERISTIC 


NUMBER 


MEAN 


STD 
DEV 


STD 
ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Richness 


9 


4.44 


1.24 


0.41 


3.49 


5.4 


Coral cover (%) 


9 


13.81 


17.74 


5.91 


0.17 


27.44 


Density (no. nr 2 ) 


9 


2.68 


1.98 


0.66 


1.15 


4.2 




Coral Species Richness 



• a-s 

• 10-11 



# 12 - 1-1 



O 0.5 1 Z 



k 




\Trpyi 




OTTSfw 

• 


177'ilVl 1 

• 








* 


Percent Coral Cover 


• 


f — ~ — ~"< 


' 


• 1.0-5.5 

• 5.6-12.0 

• 12.1- 26.0 

• 2&.1-JB.2 




Ar 


-»<^l 


% 3B.3-B9.6 


■. 






Tftn\or *2Q m JA 
0.5 1 Z 



















, 


Coral Density 
0.9 - 2 


o 


2.1 


3.7 




O 


38 


4.S 







4,7 


S3 




• 


5,9 


102 




Wiibt <&a m f 

0.5 1 2 




Figure 4.34. Midway Atoll coral statistics and distribution for 2006. Species richness (top left), percent coral cover (top 
right) and coral density (number m 2 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Kure Atoll 

Kure had the second lowest coral spe- Table 4.14. Descriptive statistics for corals from 2006 REA data at Kure 

cies richness with an average of six Atoll. Source: NWHI RAMP, unpub. data. 

species per transect (Table 4.14, Figure 

4.35). Coral cover was second lowest 

after Midway at 13.7%. Colony density 

was intermediate at 4.3 colonies/m 2 

and ranked fourth overall. 



CHARACTERISTIC 


NUMBER 


MEAN 


STD 
DEV 


STD 
ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Richness 


9 


6 


2.24 


0.75 


4.28 


7.72 


Coral cover (%) 


9 


13.67 


9.34 


3.11 


6.5 


20.85 


Density (no. m 2 ) 


9 


4.29 


2.54 


0.85 


2.33 


6.24 



Coral Spades 


Richness 




2-4 




• 


5-7 




e 


S-9 




• 


f - 11 




• 


12-14 


N 




<ftB\nr<4t> 


. A 


•J :.■ 


i 1 


2 

m Kilometers; 








Percent Coral Cover 


- 10-55 




■ 56-12.0 




• 12.1-26.0 




• 26.1-36,2 




38 3-09.6 


N 


L.i'i.l 
ftnlnr<40m 


A 


Q 0.5 1 2 







^ 



A 



Coral Density 




" 06 - 2JU 




9 2.1-3.7 




O 3 B - 4 6 




• 4.7-5.6 




:._,' 5.3-10-2 




Walcr <J0m 


A 


O 0.5 1 2 

r^ H= ^r^r^r H Kilwnatera 



Figure 4.35. Kure Atoll coral statistics and distribution for 2006. Species richness (top left), percent coral cover (top right) 
and coral density (number m 2 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Changes In Coral Cover Over Time From Rapid Ecological Assessment Data 
Most regions have low coral cover 
(<20%), with higher values at Maro, Li- 
sianski and French Frigate Shoals. Cor- 
al cover values determined from REA 
surveys showed the highest coral cover 
values at Maro and Lisianski though 
their magnitude (>60%) was greater 
than the values derived from the line- 
intercept method in 2004-2006 (Figure 
4.36). At each region, the coral cover 
values computed from the different sur- 
vey years (2004, 2005 and 2006) are 
highly similar, indicating little change 
overall in each region. 



Coral Settlement and Recruitment 
In fall 2001 an array of recruitment plates 
was attached to the base of CREWS 
moorings at French Frigate Shoals, 
Maro, Lisianski, Pearl and Hermes, Mid- 
way and Kure to assess larval recruit- 
ment. The plates were collected in fall 
2002 and fresh arrays were deployed 
which, in turn were collected in 2003. 
Coral recruits present on the plates 
were counted and measured with the 
use of a dissecting microscope. While 
the second cohort of plates were in the 
water for a shorter time period than the 
first cohort of recruitment plates that 
had been deployed between 2001 and 

2002 (293 versus 376 days, respective- 
ly), the number of recruits was higher in 

2003 (382) than in 2002 (Figure 4.37). 
Plates at Maro showed the highest 
number of recruits in both years. Nearly 
all juvenile recruits were from the family 
Pocilloporidae, though members of the 
families Acroporidae and Poritidae were 
also found on plates from French Frig- 
ate Shoals and Maro. 



Figure 4.36. Differences in coral cover among regions within the NWHI. 
Not all regions were surveyed in all three years. Coral cover was calculated 
from the line-intercept method at 0.5 m intervals. Data are mean and stan- 
dard error. Source: NWHI RAMP, unpublished data; map: L. Wedding. 




Figure 4.37. Density of coral recruits on plates deployed at six locations in 
the NWHI in 2002 and 2003. Source: NWHI RAMP; map: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Coral Bleaching 

Mass coral bleaching affected numerous 
shallow reefs through the NWHI in 2002 
and 2004 (Figure 4.38). In both years, 
the incidence of bleaching was greater 
at the three northern atolls (Pearl and 
Hermes, Midway and Kure) than at Li- 
sianski and farther south. At the three 
northern atolls, bleaching was most se- 
vere in shallow back reef and lagoon 
habitats. In both years, colonies in the 
genus Montipora and Pocillopora sus- 
tained the highest levels of bleaching 
(Kenyon et al., 2006; Kenyon and Brai- 
nard, 2006). In comparison, only low 
levels of bleaching were observed dur- 
ing 2006 surveys (Figure 4.38), which 
were conducted at the same time of 
year (September) as those in 2002 and 
2004. Colonies in the genus Montipora 
were again most affected by bleaching 
in 2006. 



1781N 




mw 


174-W 


17ZTN 




f 


dway 







T> 




1 4 


k Pearl & Hermes 


^r 




1 


\ FR PR BR 




. i 








1 \ FR PR BR 


x 








FR PR BR 


\ 


I 








LEsianski 
















Percent Coral Colonies 


with Bl cache cl Tissue- 








il 




1 


-£W 




1 2QC2 FR = Fore reef 
■ 2QQ4 PR ■ Fa(cfl re** 








A. 


Shooe BR = tock.«f 
N 
1 1 Monument Boundary A 

35 70 140 " 

^^K=^^^^^^H Kilometers 




















• # 







Figure 4.38. Percentage of colonies with bleached tissue within belt 
transects surveyed in 2002, 2004 and 2006. Minimal bleaching was seen at 
Gardner Pinnacles and French Frigate Shoals, which are not shown to re- 
duce the complexity of the figure. Source: NWHI RAMP; map: L. Wedding. 



In 2004, visual estimates of mortality 
and algal overgrowth of Montipora capi- 

tata and M. cf. turgescens at back reef sites at the three northern atolls conservatively exceeded 50%, with 
nearly complete mortality of surface-facing portions of colonies at numerous sites. The shallow crest of a large 
central patch reef system at Kure Atoll, previously referred to as "the coral gardens" due to its luxuriant growth 
of montiporids and pocilloporids, was heavily bleached in 2002. In 2004, only a few branches of Porites com- 
pressa remained alive and the dead coral skeletons were thickly covered in turf and macroalgae. Little change 
was seen in this reef's condition in 2006. A striking shift occurred at this location from a system dominated by 
coral in 2001 to a system dominated by algae in 2004 (Figure 4.39). 




Figure 4.39. Phase shift on a patch reef at Kure Atoll from a benthos dominated by coral to one dominated by algae after 
a bleaching event in 2002. Photos: J. Kenyon. 



Distribution of Coral Disease 

Coral disease has emerged as a serious threat to coral reefs worldwide and a major cause of reef deteriora- 
tion (Weil et al., 2006). The numbers of diseases and coral species affected, as well as the distribution of dis- 
eases, have all increased dramatically within the last decade (Porter et al., 2001; Green and Bruckner, 2000; 
Sutherland et al., 2004; Weil, 2004). Changing climatic conditions as well as local anthropogenic stressors 
have been implicated in increased disease levels but our ability to fully understand recent disease outbreaks 
is hampered by the paucity of baseline and epidemiological information on the normal disease levels in the 
ocean (Harvell et al., 1999, 2002). The NWHI is considered to be one of the last relatively pristine large coral 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



90 

80 

2 70 

OJ 

." 60 



C 40 



Por Trm Por TLS 



Por 
DTTS 



Por 
BND 



Mont 
TLS 



Mont 
Wspt 



Mont 
GA 



Aero 
WS 



Aero 
GA 



Poc 
WBD 



Figure 4.40. Frequency of occurrence of different coral diseases within the 
NWHI. Data based on 33 REA surveys conducted in 2005. Source: NWHI 
RAMP, unpub. data. 

Table 4.15. Distribution on of twelve coral diseases based on REA surveys 
conducted between 2002-2005. Source: Aeby unpub. data. 



reef ecosystems remaining in the world. 
As such, it provides the unique oppor- 
tunity to document the normal levels of 
disease in a coral reef system exposed 
to limited human influence and the po- 
tential of understanding disease dy- 
namics in response to changing climatic 
conditions. 

During a multi-agency cruise conducted 
in September 2002, disease investiga- 
tion was incorporated into the protocol 
and a characterization of coral diseases 
was initiated. In 2003, baseline coral 
disease surveys were conducted at 73 
permanent monitoring sites throughout 
the NWHI and have since been sur- 
veyed annually. Twelve disease states 
have now been documented in the four 
major genera of coral (Porites, Monti- 
pora, Pocillopora and Acropora) on the 
reefs of the NWHI (Figure 4.40). The 
distribution and frequency of occur- 
rence of the different coral diseases var- 
ied widely within the nine islands/atolls 
of the NWHI (Table 4.15; Figure 4.41). 
The most common disease is Porites 
trematodiasis (Figure 4.42) caused by 
the digenetic trematode, Podocotyloides 
stenometra (Aeby, 1998J. This disease 
is widespread (69.8% of the sites in 
2003) and is known to exclusively affect 
Porites sp. coral (Aeby, 2006). Other 
diseases are less common, such as Po- 
rites brown necrotizing disease, which 
only occurred at only 3.2% of the sites in 
2003 (Aeby, 2006). Patterns in disease 
prevalence among the coral genera 
suggest Acropora is highly susceptible 
to disease and Pocillopora appears to 
be very resistant. Acropora comprised only 2.2% of the overall coral community along transects, yet showed 
high overall prevalence of disease (Aeby, 2006). In contrast, pocilloporids are a common coral in the NWHI 
(21.1% of the overall coral community along transects) yet seldom showed signs of disease (Aeby, 2006). In 
contrast, Willis et al. (2004) found pocilloporids on the Great Barrier Reef to have the highest prevalence of 
disease among all coral families surveyed despite pocilloporids having the lowest coral cover. 

Although coral disease was found to be widespread on the reefs of the NWHI, disease prevalence (propor- 
tion of colonies affected) for most coral diseases, with the exception of Porites trematodiasis, was found to 
be low in a healthy ecosystem (2005: average prevalence = 0.43% SE ± 0.1%; n = 37 sites surveyed; Figure 
4.43). In contrast, the average prevalence of Porites trematodiasis in 2005 was 12.5% (SE ± 3.2%; n=36 sites 
surveyed). Porites trematodiasis is an unusual coral disease, in that, it is caused by a larval trematode, which 
is transmitted through the food chain and requires multiple hosts (coral, mollusk, fish) for completion of its life 
cycle (Aeby, 1998). In contrast, most coral diseases are caused by bacteria or viruses that can be transmitted 
directly (host to host) or through the water column. For Porites trematodiasis, prevalence of the disease is de- 





MMM 


FFS 


GAR 


MAR 


LAY 


LIS 


PHR 


MID 


KUR 


Por TRM 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Por TLS 




X 




X 


X 


X 


X 


X 


X 


Por DTTS 




X 




X 


X 


X 


X 


X 


X 


Por BND 












X 


X 






PorGA 








X 






X 




X 


MontWS 








X 


X 


X 


X 


X 


X 


Mont MFTL 








X 








X 


X 


Mont GA 




X 




X 


X 




X 






Acroporid WS 




X 
















Aero GA 




X 
















PocWB 














X 




X 


Poc GA 


X 


















Total # Diseases 


2 


6 


1 


7 


5 


5 


8 


5 


7 


Coral disease abbreviations: Por = Porites, Mont = Montipora, Aero =Acropo- 
ra, Poc = Pocillopora, TRM = trematodiasis, WS = white syndrome, WB = white 
band.TS = tissue loss syndrome, DTTS = discolored tissue thinning syndrome, 
BND = brown necrotising disease, GA = growth anomaly, MFTL = multifocal 
tissue loss 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Figure 4.41. Frequency of occurrence of different coral diseases within the 
NWHI. Data based on 33 REA surveys conducted in 2005. Source: Aeby; 
unpub. data; map: L. Wedding. 



pendent upon and thus reflective of the 
abundance of its three hosts. Hence it 
is not surprising to find high prevalence 
of this disease on the pristine reefs of 
the NWHI where host abundance is 
high. Similarly, Aeby (2007) examined 
the prevalence of Porites trematodiasis 
in Kaneohe Bay, Oahu which has a dis- 
tinct north-south gradient in rural to ur- 
banized watersheds. Most of the human 
population in surrounding watersheds is 
concentrated in the southern sector and 
so the south Bay has been most affect- 
ed by impacts of urbanization (Maragos 
et al., 1985; Hunter and Evans, 1995). 
Aeby (unpub. data) found disease lev- 
els to be higher in the north Bay than in 
the south Bay and concluded that host 
abundance, not environmental stres- 
sors, was the primary factor affecting 
prevalence of Porites trematodiasis in 
Kaneohe Bay. Prevalence of disease 
varied among islands (Figure 4.44) and 
differed among coral genera at each 
island. In 2005, Maro had the high- 
est prevalence of montiporid disease 
whereas Kure had the highest level of 
poritid disease. It is not yet clear why 
differences in disease prevalence occur 
among islands. Understanding patterns 
of disease occurrence will require much 
more information on disease epizootiol- 
ogy such as etiology and mode of trans- 
mission. 

Prevalence of the disease is depen- 
dent upon and thus reflective of the 
abundance of its three hosts. Hence it 
is not surprising to find high prevalence 
of this disease on the pristine reefs of 
the NWHI where host abundance would 

also be high. One exception to the pattern of healthy levels of coral disease on the reefs of the NWHI is an out- 
break of A cropora white syndrome (AWS) that was first documented at French Frigate Shoals in 2003 (Aeby, 
2006b). AWS has caused significant coral loss on reefs at French Frigate Shoals (Aeby, unpub. data), as well 
as in other areas of the Indo-Pacific (Willis et al., 2004; Jacobson, 2006). As of 2006, studies found the disease 
to have spread to seven reefs within French Frigate Shoals. Acroporids have also been greatly affected by 
disease in Australia (Willis et al., 2004) and the Marshall Islands (Jacobson, 2006) and have been decimated 
by disease in the Caribbean (Green and Bruckner, 2000; Porter et al., 2001; Patterson et al., 2004; Weil, 
2004). Acroporids were one of the major frame-building corals in the Florida Keys, but losses of acroporids 
are now averaging 87% or greater (Miller et al., 2002; Patterson et al., 2002). Research is desperately needed 
to understand disease processes to prevent the acroporid reefs of the Monument and neighboring Johnston 
Atoll National Wildlife Refuge from following the same path as the acroporid reefs of the Florida Keys. More 
research is also needed to understand disease transmission dynamics across Indo-Pacific and the impact 
climate change will have on the health of coral reef ecosystems. 




Figure 4.42. Porites trematodiasis is caused by a larval trematode, which is 
transmitted through the food chain and requires multiple hosts (coral, mol- 
lusk, fish) for completion of its lifecycle. Photo: G. Aeby 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



p- 


French Fi 
Shoals 


igate - | 

1 ^ 


l 


Coral Disease 
(Presence/Absence) 

J 

□ Ac,„po,, 

□ L„« N 

~ A 

2 4 
^^■zzzzzn Kilomeiers 




L 1 



Maro Reef 



Jl 



Coral D 


sease 


(Presence/Abse 


nee) 


J 






A,-,,„ 


ra 




^H Porite 


Trema 


ud. 


^H Porile 


other 




^| Monti 






□ uw 




N 




•:20 m 


A 










zi Kilometers 




Lisianski 



Coral Disease 
(Presence/Absence) 



«„„„.. 






1.5 3 



A 



173*48™ 176=6^ 




168°0'20 M W 



167"59'40 , 'W 



Gardner Pinnacles 



Coral Disease 


(Presence/Absence) 


J 




| | A=,»po„ 




^^ Pontes Tremalod. 


^H Porttes other 




^| Montipora 




□ l.h. 


N 

A 


Water <20 m 


0.1 0.2 










Laysan 






^ 








Coral Disease 
(Presence/Absence) 

J 




w -" !0 ™ A 






^^^m i Kilometers 









Pearl and 

Hermes J. 


J 


lit 






1^ 
" 1" 

™*" ^(i!P» CfTS 


1 


Coral Disease 
(Presence/Absence) 

Acropora 


Water <20 m J\ 
2 4 

^^^■zzzzzzi Kilometers 


\ 





Figure 4.43. Coral disease for French Frigate Shoals, Gardner Pinnacles, Maro Reef, Laysan, Lisianski and Pearl and 
Hermes. Source: Aeby, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Midway Atoll 




Coral Disease 
{Presence/Absence) 



| Acropor 

^| Pomes Ti 
^H Pontes ol 



Kure Atoll 



.11 



Coral Disease 


(Presence/Absence) 


A 




,,„„ 




^| Porites 1 1 em 


tod. 


^H pontes other 




^| Montlpora 




^L,„d 


N 

A 


Water <20 m 






^™=i Kilometer 





1 

1 




Figure 4.43 (continued). Coral disease at Midway and Kure atolls. Source: Aeby, unpub. data; maps: L. Wedding. 





1.0 -, 




0.030 -, 








0.9 - 




UAcropora 


□ Pocillopora 






0.8 - 








0.025 - 










0.7 - 








0.020 - 










0.6 - 


















0.5 - 












0.015 - 










0.4 - 












0.010 - 














0.3 - 


























0.2 - 












0.005 - 












CD 


0.1 - 
0.0 - 










0.000 - 














i i i i 




i i i 


1 




O 


FFS MAR 


PHR MID KUR 


FFS MAR 


PHR MID KUR 


CD 










> 
CD 


50 -I 




1.0 - 






Q. 


45 - 


□ Pontes 




0.9 - 


□ Montipora 






40 - 








0.8 - 










35 - 








0.7 - 










30 - 








0.6 - 










25 - 










0.5 - 












20 - 










0.4 - 












15 - 
10 - 




rf 




4- 








0.3 - 
0.2 - 












5 - 










r+i 




0.1 - 




























m 
















r+n 










FFS MAR 


PHR MID KUR 


i 0.0 i 

FFS MAR 


PHR MID KUR 



Figure 4.44. Prevalence of coral disease among locations by major coral genera. Note the differences in y-axis among 
genera. Source: G. Aeby unpub. data. 



Coral Predators 

The Crown-of-thorns starfish (COTS; Acanthaster planci; Figure 4.45) and Drupellid snails (Drupella sp.) 
are both corallivores that have caused significant coral damage in other areas of the Indo-Pacific, and were 
monitored on the reefs of the NWHI during benthic (COTS and Drupellid snails) and towed-diver surveys 
(COTS). Towed-diver surveys report COTS to be present on the reefs of the NWHI but to occur at low levels 
(average=0.65 COTS/km; NWHI Reef Assessment and Monitoring Program or RAMP, unpub. data). During an- 
nual benthic monitoring surveys it was found that the frequency of occurrence (the number of sites with animals 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



or feeding scars/total sites surveyed) for 
Drupella sp. was low (3% in 2004, 15.4% 
in 2005) (Aeby, unpub. data). Drupella 
were usually found feeding at the base 
of branches of cauliflower coral (Pocil- 
lopora meandrina). For COTS, frequen- 
cy of occurrence was also low with re- 
ports of COTS observed at 4.5% of the 
monitoring sites in 2004 and 28.2% in 
2005 (Aeby, unpub. data). COTS were 
usually found as single animals. 




Figure 4.45. Crown-of-thorns sea star in the NWHI. Photo: J. Kenyon. 



ALGAE 

Although the NWHI represent one of the last relatively intact tropical reef ecosystems in the world, macroal- 
gal community dynamics of the 10 atolls, islands, and reefs situated in the NWHI Marine National Monument 
remain poorly understood. A study published in conjunction with the Northwestern Hawaiian Islands' third Sci- 
entific Symposium (Vroom and Page, 2006) was the first to provide distributional maps of common algal spe- 
cies, statistically compare sites from differing habitats and islands based on relative abundance of macroalgae 
and look for temporal differences in macroalgal populations. Findings revealed that the abundance of most 
macroalgal genera was low across the archipelago, but that members of certain green algal genera including 
Halimeda and Microdictyon (Figure 4.46) can be extremely common and in some cases form dense monotypic 
meadows on the reef, especially in fore reef areas (Microdictyon) and lagoons (Halimeda). Other genera, such 
as the brown algae Stypopodium and Lobophora, and the red alga Laurencia, become increasingly prevalent 
in the three northwestern-most atolls of the Hawaiian archipelago (Kure, Midway, and Pearl and Hermes). 
Relative abundance of macroalgae across the NWHI chain as a whole remained relatively static for the years 
surveyed; however, slight changes occurred at Kure and Midway atolls where coral bleaching events were 
documented in 2002 and 2004. 



-~ 


1V3? -;^^E f?3pSfi%%i 1 




^fcJ<rir I HSi^Br.^B 


S9^^' K^^^fl ItW 


, _^r * Hi 


■&?££> ** ■ 




^ 3Q^BSESSKP?3Bfe'^ii. "HP! 










*<£ --if- V-^-ife-? j 


« & \4z - - 




M - ^ 




y 


M- " M 






3k. ^e-7.- {■?, ■l^rt <■ 










Figure 4.46. Halimeda velasquezii at Maro Reef (left). H. velasquezii Is the most common species of Halimeda in the 
NWHI. Microdictyon is also very common in some regions of the NWHI. A close-up photo of a Microdictyon setchellianum 
(right) field at Gardner Pinnacles. Photos: P. Vroom. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

A study recently completed at Pearl and Hermes Atoll (Page, 2006) used detailed species-level percent cover 
analyses coupled with environmental variables to better understand the mechanisms that determine distribu- 
tional patterns of organisms, particularly algae. Benthic community composition was examined along a wave 
exposure gradient using multivariate statistical analyses with the hypothesis that sites with similar levels of 
wave exposure would exhibit similar benthic communities. Species richness of coral and macroalgae were 
also compared to determine if sites with intermediate levels of wave exposure would contain the highest di- 
versity of benthic organisms. To test these hypotheses, percent cover of benthic organisms was determined 
at 34 sites in four wave exposure categories: high, intermediate-high, intermediate-low and low. Multivariate 
statistical analyses revealed that sites from each wave exposure category differed significantly, and a non- 
metric multi-dimensional scaling ordination (nMDS) and cluster diagram grouped sites from low, high, and 
intermediate-high wave disturbance areas into three relatively discrete clusters. However, sites experiencing 
intermediate-low wave exposure did not group together in the nMDS ordination or cluster diagram, suggesting 
variability in benthic compositions among these sites. Coral and macroalgal species richness was significantly 
higher at sites with intermediate-high and intermediate-low levels of wave exposure than at sites with low wave 
exposure, although not significantly higher than sites with high wave exposure. 

An article appearing in American Scientist magazine (Vroom et al., 2006) compared percent cover of macroal- 
gal, turf algal, crustose coralline algal and coral populations at eight islands across the Pacific Ocean basin, 
including two from the NWHI. The NWHI are documented to contain the highest percent cover of algal species 
when compared to other geographic locations, and the lowest percent cover of living coral. This is likely due 
to the subtropical location of the NWHI and cool SSTs that bathe biological communities during winter months. 
Despite high algal populations, the NWHI remain healthy and thriving marine ecosystems that are dominated 
by top predators and high fish populations. 



Algal diversity appears similar across 
the NWHI chain even though brown al- 
gae tend to be more abundant at Mid- 
way and Kure atolls when compared to 
most other islands (Figure 4.47). The 
lower abundance of green algae at Mid- 
way may be tied to lower apex preda- 
tor biomass and higher herbivorous fish 
densities at this atoll system, suggesting 
possible top-down control of the ben- 
thic habitat (DeMartini and Friedlander, 
2004, 2006). 

Although the mix of macroalgal species 
is relatively similar throughout the NWHI 
chain, certain species (e.g., Stypopo- 
dium flabelliforme, Laurencia galtsoffii) 
are more abundant in the northwestern- 
most atolls where SSTs experience the 
greatest annual fluctuation. While S. 
flabelliforme is a major component of 
shallow reef systems at Kure Atoll, it is a 
minor component of reefs at most other 

islands and atolls in the NWHI. Because brown algae are known to predominate over other algal lineages in 
cool, temperate environments (Cheney, 1977), it is possible that the cooler SSTs found at Kure and Midway 
atolls during winter months may favor a higher abundance of brown algal species (Figure 4.47). 




Figure 4.47. Prevalence of major algal lineages in the Northwestern Ha- 
waiian Islands. Bars represent standard deviation. Source: NWHI RAMP, 
unpub. data; map: L. Wedding. 




an 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Algae Data: 2006 

Mokumanamana 

Only two stations were assessed for 
algae at Mokumanamana in 2006 (Fig- 
ure 4.48; located on pages 144-145). It 
is a composite of maps illustrating the 
percent frequency of occurrence of ma- 
jor algae by island). Four algal groups 
were present with Halimeda occurring in 
100% of the samples, followed by Lau- 
rencia (91.7%), crustose coralline red 
algae (12.5%) and Jania (8.3%; Table 
4.16). 



Table 4.16. Summary statistics for algal groups at Mokumanamana Island 
in 2006. Source: NWHI RAMP, unpub. data. 


STATIASTICS 


MEAN 


SD 


FREQ 


% FREQ 


Dictyosphaeria 











0.00% 


Halimeda 


100 





2 


100.00% 


Microdictyon 











0.00% 


Neomeris 











0.00% 


Jania 


8.33 


11.79 


1 


50.00% 


Laurencia 


91.67 


11.79 


2 


100.00% 


Non-geniculate branched coralline red 
algae 











0.00% 


Crustose coralline red algae 


12.5 


17.68 


1 


50.00% 


Lobophora 











0.00% 


Padina 











0.00% 



French Frigate Shoals 
Ten stations were sampled in 2006 at 
French Frigate Shoals with seven algal 
groups present on quantitative surveys 
(Table 4.17, Figure 4.48). Laurencia 
was present all stations and found, on 
average, in 77% of quadrats. Halimeda 
was present at 90% of the stations and 
found, on average, in 62% of the quad- 
rats. Jania was found at half the stations 
but on occurred in 9% of the quadrats, 
on average. 



Table 4.17. Summary statistics for algal groups at French Frigate Shoals 
in 2006. Source: NWHI RAMP, unpub. data. 



STATIASTICS 


MEAN 


SD 


FREQ 


% FREQ 


Dictyosphaeria 


1.67 


5.27 


1 


10.00% 


Halimeda 


62.5 


38.14 


9 


90.00% 


Microdictyon 


15.83 


21.68 


4 


40.00% 


Neomeris 











0.00% 


Jania 


9.17 


15.44 


5 


50.00% 


Laurencia 


76.67 


20.71 


10 


100.00% 


Non-geniculate branched coralline red algae 











0.00% 


Crustose coralline red algae 











0.00% 


Lobophora 


30 


32.68 


8 


80.00% 


Padina 


2.5 


7.91 


1 


10.00% 



Maro Reef 

Nine stations were sampled at Maro 
Reef in 2006 with six algal groups pres- 
ent (Table 4.18, Figure 4.48). Halimeda 
was present at all stations and found, on 
average, in 91% on the quadrats. Lau- 
rencia was also found at all stations and 
occurred in 75% of the quadrats. Crus- 
tose coralline red algae were found at 
three-quarters of the stations surveyed 
and appeared an average of 34% of the 
quadrats. 



Table 4.18. Summary statistics for algal groups at Maro Reef in 2006. 
Source: NWHI RAMP, unpub. data. 



STATIASTICS 


MEAN 


SD 


FREQ 


% FREQ 


Dictyosphaeria 


5.56 


8.33 


4 


44.44% 


Halimeda 


90.74 


6.51 


9 


100.00% 


Microdictyon 











0.00% 


Neomeris 











0.00% 


Jania 


12.96 


19.59 


4 


44.44% 


Laurencia 


75 


19.09 


9 


100.00% 


Non-geniculate branched coralline red 
algae 











0.00% 


Crustose coralline red algae 


34.26 


36.43 


6 


66.67% 


Lobophora 


27.78 


30.33 


6 


66.67% 


Padina 











0.00% 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Laysan Island 

Three stations were surveyed at Lasyan 
Island in 2006 with eight algal groups 
observed in these surveys (Table 4.19, 
Figure 4.48). Halimeda and Laurencia 
were found at all stations and were the 
dominant groups being found, on aver- 
age, in 89% and 86% of all quadrats, re- 
spectively. Crustose coralline red algae 
occurred at two of the three stations and 
were present in 44% of the quadrats, on 
average. 



Table 4.19. Summary statistics for algal groups at Laysan Island in 2006. 
Source: NWHI RAMP, unpub. data. 



Lisianski-Neva Shoals 
Eight stations were sampled at Lisians- 
ki-Neva Shoals in 2006 with eight algal 
groups present in those surveys (Table 
4.20, Figure 4.48). Halimeda occurred 
at all stations and in nearly all quadrats 
(94%). Other important genera included 
Laurenica, which was present in 78% of 
the quadrats, and Jania, which occurred 
in 42% of the quadrats. Both genera 
were present at 89% of the stations sur- 
veyed. 



Pearl and Hermes Atoll 
A total of 12 stations were sampled at 
Pearl and Hermes in 2006 with nine 
algal groups present in those surveys 
(Table 4.21; Figure 4.48). Laurencia oc- 
curred at the greatest number of stations 
(92%), followed by Halimeda (75%), 
Microdictyon (67%), Dictyosphaeria 
(67%) and Lobophora (58%). On aver- 
age Halimeda was present in 49% of 
quadrats, followed by Microdictyon with 
42%, and Laurencia with 40%. Although 
Dictyosphaeria had a high frequency of 
occurrence, it was only present in 12% 
of the quadrats, on average. 



STATIASTICS 


MEAN 


SD 


FREQ 


% FREQ 


Dictyosphaeria 











0.00% 


Halimeda 


88.89 


19.25 


3 


100.00% 


Microdictyon 


19.44 


33.68 


1 


33.33% 


Neomeris 


2.78 


4.81 


1 


33.33% 


Jania 


8.33 


14.43 


1 


33.33% 


Laurencia 


86.11 


17.35 


3 


100.00% 


Non-geniculate branched coralline red 
algae 











0.00% 


Crustose coralline red algae 


44.44 


38.49 


2 


66.67% 


Lobophora 


19.44 


4.81 


3 


100.00% 


Padina 


5.56 


9.62 


1 


33.33% 



Table 4.20. Summary statistics for algal groups at Lisianski-Neva Shoals 
in 2006. Source: NWHI RAMP, unpub. data. 


STATIASTICS 


MEAN 


SD 


FREQ 


% FREQ 


Dictyosphaeria 


12.96 


13.89 


6 


66.67% 


Halimeda 


93.52 


8.1 


9 


100.00% 


Microdictyon 


25 


33.59 


5 


55.56% 


Neomeris 


10.19 


11.62 


5 


55.56% 


Jania 


42.59 


33.19 


8 


88.89% 


Laurencia 


77.78 


31.18 


8 


88.89% 


Non-geniculate branched coralline red 
algae 











0.00% 


Crustose coralline red algae 


1.85 


5.56 


1 


11.11% 


Lobophora 


25 


16.14 


8 


88.89% 



Table 4.21. Summary statistics for algal groups at Pearl and Hermes Atoll 
in 2006. Source: NWHI RAMP, unpub. data. 



STATIASTICS 


MEAN 


SD 


FREQ 


% FREQ 


Dictyosphaeria 


11.81 


12.54 


8 


66.67% 


Halimeda 


49.31 


39.64 


9 


75.00% 


Microdictyon 


41.67 


44.1 


8 


66.67% 


Neomeris 


4.86 


10.33 


3 


25.00% 


Jania 


20.14 


23.15 


6 


50.00% 


Laurencia 


40.28 


31.15 


11 


91.67% 


Non-geniculate branched coralline red 
algae 











0.00% 


Crustose coralline red algae 


18.75 


32.4 


4 


33.33% 


Lobophora 


29.86 


36.66 


7 


58.33% 


Padina 


10.42 


26.38 


3 


25.00% 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Midway Atoll 

Sampling at Midway Atoll in 2006 includ- 
ed six stations with eight algal groups 
present (Table 4.22, Figure 4.48). Lau- 
rencia was found at all stations and 39% 
of the quadrats. Lobophora was found 
at 83% of the stations and occurred in 
47% of the quadrats, on average. Mic- 
rodicyton was found at half the stations 
and occurred in 22% of the quadrats. 
Padina was present at two-thirds of the 
stations (19% of the quadrats). 



Ku re Atoll 

Eight stations were sampled at Kure 
Atoll in 2006 with eight algal groups 
present (Table 4.23, Figure 4.48). Mic- 
rodictyon was the most abundant gen- 
era, occurring at all stations and in 62% 
of all quadrats. Halimeda also was pres- 
ent at all stations but in a lower percent- 
age of quadrats (25%). Jania and Lobo- 
phora were both present at 87% of the 
stations. 



Table 4.22. Summary statistics for algal groups at Midway Atoll in 2006. 
Source: NWHI RAMP, unpub. data. 

STATISTICS MEAN SD FREQ % FREQ 



Dictyosphaeria 


6.94 


13.35 


2 


33.33% 


Halimeda 


11.11 


16.39 


3 


50.00% 


Microdictyon 


22.22 


32.77 


3 


50.00% 


Neomeris 











0.00% 


Jania 


1.39 


3.4 


1 


16.67% 


Laurencia 


38.89 


19.48 


6 


100.00% 


Non-geniculate branched coralline red algae 











0.00% 


Crustose coralline red algae 


12.5 


16.46 


3 


50.00% 


Lobophora 


47.22 


24.53 


5 


83.33% 



Padina 



19.44 



21.52 



66.67% 



Table 4.23. Summary statistics for algal groups at Kure Atoll in 2006. 
Source: NWHI RAMP, unpub. data. 


STATISTICS 


MEAN 


SD 


FREQ 


% FREQ 


Dictyosphaeria 


10.42 


19.8 


4 


50.00% 


Halimeda 


25 


23.15 


8 


100.00% 


Microdictyon 


62.5 


26.35 


8 


100.00% 


Neomeris 











0.00% 


Jania 


20.83 


15.43 


7 


87.50% 


Laurencia 


43.75 


37.73 


6 


75.00% 


Non-geniculate branched coralline red algae 











0.00% 


Crustose coralline red algae 


31.25 


36.39 


4 


50.00% 


Lobophora 


30.21 


27.44 


7 


87.50% 


Padina 


6.25 


11.57 


3 


37.50% 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Mokumanamana 

JJL 



*& Freq. ol Algae 




z IGBPZVW 



iiL 

— ^ 

AL IJl 



French Frigate 
Shoals 



% Freq. of Algae 



□ >- 




Maro Reef 



% Freq. of Algae 



^| Microdictyon 



Lobciptioru 
^| Padina 



<20m f\ 




Laysan Island 



% Freq. of Algae 




Lisianski 


1L1L Q ^% 


% Freq. of Algae 

^H Dlctyosphaeria 

ZJ M,c,„d lU »„„ 

^| Padina 

□ -d N 

Water <20 m f\ 
12 4 





Pearl and 
Hermes 



Lll 

JJiL 



% Freq. of Algae 

^| DiclyospDaeria 

Jai'.ia 



□ ■• 



A 



JUJU 






IiL 



ii-JkL 



Figure 4.48. Percent frequency of occurrence of major algal groups by island in 2006. Source: NWHI RAMP, unpub. data; 
maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



177°2T\N 


177°24'W 


177°2rW 


Midway 


"* 




%Freq. of Algae 






H r ,, ,, ,,.„. 






"m"™ 


JliU 




■ »nl. 


f~~~y 


* L- 11 


5. u ~-». 




5 ><1) 1 


^| Pad ina 
o 0.5 i a 


-M^dx 


u 





Kure 



% Freq. of Algae 




-1 




^| Dictyospriaeria 




^H Halimeda 




Microdiclyon 




^B Neameris 




ED j™ 




^H umrencta 




^H crustose coral 


aa 


^H Labaphara 




^| Padina 


N 


Water <40 m 


A 






^■=^^™Kilc 





MiJJIl 



JiiM 



iL 



M 




Figure 4.48 (continued). Percent frequency of occurrence of major algal groups by island in 2006. Source: NWHI RAMP, 
unpub. data; maps: L. Wedding. 



INVERTEBRATES 

Census of Coral Reef Ecosystems (CReefs) 

The international CoML is a global effort to assess the diversity, distribution and abundance of ocean life and 
explain how it changes over time. Over 1,700 scientists from 73 countries are pooling their findings to cre- 
ate a comprehensive and authoritative portrait of life in the oceans today, yesterday and tomorrow. As one 
of 17 projects of the CoML, the goals of the CReefs are to increase tropical taxonomic expertise, conduct a 
taxonomically-diversified global census of coral reef ecosystems, and improve access to and unify coral reef 
ecosystem information scattered throughout the world. 

As part of the CReefs effort, NOAA's Pacific Islands Fisheries Science Center Coral Reef Ecosystem Division 
led a multi-institutional team of international taxonomists on a 23-day research expedition in October 2006 to 
explore the biodiversity of small, understudied, or lesser known invertebrate, algal and microbial species at 
French Frigate Shoals. In an effort to maximize the ability to document biodiversity, surveys were conducted 
at over 50 different sites representing 14 habitat types using 12 diverse sampling methods (including baited 
traps, rubble brushing, rubble extraction, underwater vacuuming with gentle suction, plankton tows, light traps, 
sediment and water sampling) specifically designed to minimize habitat impacts while maximizing the number 
of ecological niches sampled. 

During the three week cruise, scientists documented more than 1,000 species at French Frigate Shoals. For 
comparison, this corresponds to around 20% of the Hawaiian marine invertebrate fauna documented over 
the past 200 years. These new findings indicate just how little is known about tropical marine invertebrates in 
general and especially within the NWHI, and may offer clues to the extent of undiscovered diversity at French 
Frigate Shoals. Collected species were photo-documented for future study. Many species had never been 
photographed, fresh or alive, and thus represent the first documentation of their living color and appearance. 
DNA samples were also collected to facilitate DNA-based identification of Hawaiian and tropical Pacific inver- 
tebrates in the future. 



Thorough taxonomic identifications and molecular analyses of the samples collected are still being analyzed 
and will take many years to complete. However, preliminary findings suggested that approximately 2,300 
unique morpho-species were collected and photographed during the 16 days of sampling (Figure 4.50). To 
improve the long-term ability to monitor biodiversity, tissue samples for molecular barcoding were collected 
from about 60% of the unique morpho-species. An estimated 30-50 collected specimens are thought to be new 
species to science, including new species of crabs, corals, sea cucumbers, sea quirts, worms, sea stars, snails 
and clams. From this expedition, well over a hundred new species records from sponges, corals, anemones, 
flatworms, segmented worms, hermit crabs, crabs, sea slugs, bivalves, gastropods, octopus, sea cucumbers, 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

sea stars and sea squirts, will likely be 
identified for French Frigate Shoals. The 
highest sampled diversities at French 
Frigate Shoals were in the phylums An- 
thropoda and Mollusca. By habitat type, 
lagoon patch reefs, La Perouse Pinnacle 
(basalt), back reef and deep fore reefs 
had the highest diversity. Due to the 
high level of taxonomic expertise avail- 
able on the cruise, hand collection was 
the most effective sampling methodolo- 
gy, following by rubble extraction, rubble 



800 -, 
700 
600 
500 - 
400 - 
300 
200 - 
100 - 




I 



■J 



a 



d_ 



*p & 



«sr 



#■ 









$> 
J 



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# .*# ^ & <s^ 



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brushing and use of baited traps. 

Some anecdotal figures indicate the de- 
gree of novelty. For example, of the six 
specimens of octopus collected three 
species may be new to science (Figure 
4.50). In comparison, 15 species of oc- 
topus were previously known from the Hawaiian Islands. Additionally, there were at least 25 species of sea 
cucumbers documented; three are new species and two are new records for Hawaiian fauna (Figure 4.50). 
Previously, about 30 species of shallow water holothurians have been documented from Hawaiian waters. 



Figure 4.49. Unique morphospecies collected at French Frigate Shoals by 
phylum from CReefs cruise. Source: CoML, unpub. data. 




Figure 4.50. Six species of octopus (left) were collected during the mission, three of which maybe new to science. The 
right photo shows a newly discovered species of sea cucumber. Photos: G. Paulay 

Though relatively high diversity was found for sponges, bryozoans, eulimid gastropods, hermit crabs, echino- 
derms, ascidians and other invertebrates (including corallimorph anemones, galatheid squat lobsters, porcel- 
lanid crabs, pea crabs and coral barnacles), had strikingly low diversity or were absent. Interestingly, about 
one third of all invertebrate morphospecies collected were either found only once or found at only one site. A 
possible new family of ascidian for the NWHI, Mogulidae, was collected. Likewise, a new species of coral that 
could not even be identified to family level was found and photographed, since coral collections were not au- 
thorized under this permit. An estimated 48 new species records of Opisthobranch molluscs for French Frigate 
Shoals were collected, 27 of which appear to be new records for the NWHI. 



Other Invertebrate Surveys 

Recent efforts to quantify the non-coral invertebrate populations in the NWHI included two broad-scale towed- 
diver surveys conducted in 2004 and 2006 (Figure 4.51) and a REA survey conducted in 2005. Surveys were 
focused on collecting information on three target classes of invertebrates: Echinoidea, Holothuroidea and As- 
teroidea. Towed-diver surveys found densities of echinoids and holothuroids to be highest at the northernmost 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 






Figure 4.51 Mean density of Echinoids, Asteriods and Holothuroids per m 2 from towed diver surveys. Sources: NWHI 
RAMP, Brainard et a/., in prep.; maps: L. Wedding. 

islands/atolls. Sea urchins were the most common invertebrate observed during these surveys, with Kure 
(2004 and 2006) and Midway (2006) reporting the highest densities in the island chain (>1,600 urchins/ha). 
Sea cucumbers were present at all islands but in low densities, with the exception of the northern atolls. The 
highest sea cucumber density was recorded at Kure in 2006. COTS were in relatively low abundance through- 
out the archipelago with the highest density recorded in 2004 at Pearl and Hermes. Though abundance of 
COTS was relatively low in comparison with reported infestation levels in higher coral cover areas, such as the 
Great Barrier Reef, the impacts of these abundances could be significant due to the relatively low coral cover 
found throughout much of the NWHI. 

Data collected during REA surveys included species level information on the three target classes of inverte- 
brates and followed the general patterns of the towed-diver data. The most common echinoid throughout the 
NWHI was the burrowing sea urchin, Echinostrephus sp., with the highest densities recorded at Midway and 
Kure (>12 individuals/m 2 ). As in towed-diver surveys, sea cucumbers were present at all islands/atolls but in 
low densities. The most common sea cucumber was Actinopyga obesa, with a density of .03 individuals/ m 2 at 
Kure. The most common sea-star was Linckia multifora. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

EXISTING DATA GAPS 

Effective management decisions are based on reliable information on the biological characteristics of the or- 
ganisms, their ecological relationships and understanding of the natural temporal variations that characterize 
their ecosystems. Overall there is a need for basic information on all living resources at the NWHI. Taxonomy 
studies should provide knowledge of all species presents to develop a baseline and facilitate identification of 
new species records and which ones are endemic, rare and worthy of special attention. Life history studies 
are needed to provide information on essential habitat requirements (reproduction, recruitment and feeding) 
for all life stages, environmental tolerance, larval dispersal mechanisms and other parameters (age structure, 
growth, mortality, etc.) for key native species. Specific research opportunities include: 

• Developing a comprehensive catalog of native species that include: ecological requirements, georef- 
erenced habitat use during different life stages, life history metrics (population age/size distributions, 
growth rates, size/age of maturity, mortality rates, etc.); 

• Comparing native species habitats and ecological requirements to a catalog of anthropogenic threats in 
order to identify native species vulnerable to anthropogenic stress; 

• Developing comparative life history studies between NWHI and MHI to determine anthropogenic effects 
on growth, maturation, and reproductive success of native species; 

• Documenting the trophic dynamics of key native species; 

• Developing a comprehensive species list of algal taxa for the NWHI; 

• Determining habitat distribution/availability, especially that essential to reproduction, recruitment, and 
feeding; 

• Characterizing the genetic structure of specific populations; and 

• Determining the functional ecological roles of native species. 

Monitoring programs require research to determine priority parameters and indicators to be measured for 
changes to ecosystems and ecological processes. Research also needs to indicate the scale, resolution and 
frequency of sampling that will contribute to meaningful monitoring, and the kinds of tools and technology that 
can best obtain the desired data. There are key gaps in the development and implementation of monitoring 
programs, including the need to determine: 

• The variables, scale and the spatial and temporal resolution at which ecological processes and connec- 
tivity can be monitored to support management needs; 

• The existing parameters and indicators of existing monitoring programs and identify gaps as the basis 
for more comprehensive monitoring of ecosystem change; and 

• The spatial and temporal basis of ecological processes to identify ecological boundaries between sub- 
regions. 

There are also key research issues that have been identified as important for monitoring and modeling, includ- 
ing: 

• Which environmental conditions, e.g., temperature, flow, geomorphology, have a mitigating influence on 
survival in a changed environment; 

• To what extent the reduction or expansion of one or more segments of the community assemblage result 
in competitive top down pressure or an increase in bottom up production; 

• How ecosystem acclimation to change varies among taxa and in relation to both survival and the ability 
to effectively reproduce; 

• The degree to which variability in an ecosystem may determine its capacity for resilience; 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

• How the rebound of an ecosystem depends on maintenance of established pathways of energy flow, 
which provide the system a stable means of recovery rather than risk a transition to a different state of 
equilibrium; 

• To what extent reducing fish populations of the ecosystem undermine or realign energy flow and trophic 
stability; 

• The other key aspects that affect ecosystem stability and resilience need to be identified, e.g., rates of 
energy flow, oceanographic conditions, nutrient levels and recruitment; 

• The spatial and temporal patterns of plankton and larval dispersal, sources and sinks and 

• The size, location and effectiveness of closed areas, e.g., MPAs. 

As with monitoring of ecosystems and ecological processes, there is a need for research on the parameters 
and indicators most appropriate and practical for measuring changes to species populations and habitats. Re- 
search needs to address both marine terrestrial biodiversity and communities and seek to identify the scale, 
resolution and frequency of sampling that will contribute to meaningful monitoring of flora and fauna, as well 
as the most useful tools and technology for gathering the data. Important gaps in the development and imple- 
mentation of monitoring programs include the need to determine: 

• The variables, scale and the spatial and temporal resolution at which biodiversity and habitats should be 
monitored to support management needs; 

• The existing parameters and indicators of existing monitoring programs and identify gaps as the basis 
for more comprehensive monitoring; and 

• What spatial and temporal parameters distinguish sub-populations of species or differentiate habitats in 
the NWHI. 

There are also specific research issues that have been identified as important for monitoring biodiversity and 
habitats, including the need to: 

• Determine the priority species for monitoring in relation to anticipated short-term impacts, including from 
management actions, and long-term change; 

• Document 'hot spots' of adult population abundance; 

• Determine the movement of key species into the NWHI and within the NWHI; 

• Assess priority species populations as a basis for monitoring and developing recovery models and pro- 
jections of future population levels; 

• Undertake life history studies for all priority species to provide information on essential habitat require- 
ments (reproduction, recruitment and feeding) for all life stages; 

• Determine essential habitat and ecological requirements of protected species to minimize anthropogenic 
threats and the effect of catastrophic events; 

• Determine which likely effects of climate change on protected species are priorities for monitoring, e.g., 
the effect of sea level rise on nesting site of the green sea turtle and Hawaiian monk seal; and 

• Evaluate exiting and potential diseases affecting priority species and habitats of the NHWI and develop 
appropriate methods to monitor the presence and impact of these on terrestrial and marine biodiversity. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



APPENDIX 

Distribution of corals and anemones reported in the NWHI during 1907-2006 compiled by Maragos from 
Vaughan (1907), Dana (1971), Maragos et al. (2004), and the unpublished records of G. Aeby, J. Asher, 
Maragos, B. Vargas and B. Zgliczynski. Endemics in bold and asterisks (*) indicate undescribed species. 



Dana (1846), 
J. Kenyon, J. 




STONY CORALS 


NIH 


NEC 


FFS 


GAR 


MAR 


LAY 


LIS 


PHR 


MID 


KUR 


N 


*Coral unid., seen first by J. Starmer, sp.18 






X 
















1 


Acropora cerealis 






X 


X 


X 












3 


A. cytherea 




X 




X 


X 


X 




X 






5 


A. gemmifera 






X 


X 














2 


A. humilis 






X 


X 


X 












3 


A. nasuta 






X 




X 


X 










3 


A. paniculata 






X 
















1 


*A. sp.l (prostrate) 






X 








X 








2 


*A. sp.28 cf. retusa 






X 
















1 


A. valida 






X 




X 


X 


X 


X 






5 


*A. sp.29 (table) 






X 
















1 


*A. sp.30 cf. palmerae 






X 
















1 


A. sp. 20 (neoplasia/tumor?) 






X 
















1 


A. sp.26 cf. loripes 






X 
















1 


Montipora capitata 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


11 


M. flabellata 




X 


X 


X 


X 


X 


X 


X 


X 


X 


9 


M. patula 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


10 


*M. sp.4 cf. incrassata 




X 


X 




X 










X 


4 


M. dilatata 












X 


X 








2 


*M. sp.6 cf. dilatata 










X 












1 


*M. sp.7 (foliaceous) 






X 








X 


X 


X 




4 


*M. sp.2 (ridges) 
















X 




X 


2 


*M. sp.5 (branching) 














X 








1 


*M. sp.14 (nodular) first seen by B. Vargas 
















X 






1 


M. tuberculosa 






X 




X 


X 


X 


X 


X 


X 


7 


*M. sp.24 (irregular) 






X 
















1 


*M. sp.3 cf. turgescens 










X 


X 


X 


X 


X 


X 


6 


M. verrilli 






X 




X 


X 


X 


X 


X 


X 


7 


Gardineroseris planulata 


















X 




1 


Leptoseris hawaiiensis 






X 






X 










2 


L. incrustans 






X 










X 


X 


X 


4 


*L. sp.22 cf. incrustans 






X 
















1 


L. mycetoseroides 






X 
















1 


*L. cf. papyracea spl9 






X 
















1 


*L. cf. scabra spl7 






X 








X 








2 


Pavona clavus 
















X 


X 


X 


3 


P. duerdeni 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


10 


P. maldivensis 






X 




X 




X 


X 


X 


X 


6 


P. varians 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


10 


*Balanophyllia sp. (pink) 






X 




X 










X 


3 


Cladopsammia eguchii 






X 


X 


X 


X 




X 


X 


X 


7 


Tubastraea coccinea 


X 




X 


X 


X 


X 




X 


X 


X 


9 


Cyphastrea ocellina 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


10 


Leptastrea agassizi 






X 




X 








X 




3 


L. bewickensis 






X 








X 


X 






3 


L. purpurea 


X 


X 


X 


X 


X 


X 




X 


X 


X 


9 


L. pruinosa 




X 


X 


X 


X 












4 


*L. sp.8 cf. F. hawaiiensis 




X 


X 




X 




X 






X 


5 


*Cycloseris tenuis 






X 


X 






X 


X 






4 


*C. vaughani 






X 








X 


X 


X 




4 


Diaseris distorta 






X 








X 








2 





A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Continued from previous page, distribution of corals and anemones reported in the NWHI during 1907-2006 compiled 
by Maragos from Dana (1846), Vaughan (1907), Dana (1971), Maragos et al. (2004), and the unpublished records of G. 
Aeby, J. Asher, J. Kenyon, J. Maragos, B. Vargas and B. Zgliczynski. Endemics in bold and asterisks (*) indicate unde- 
scribed species. 



STONY CORALS 


NIH 


NEC 




GAR 


MAR 


LAY 




PHR 


MID 


KUR 


N 


Fungia scutaria 


X 




X 


X 


X 


X 


X 


X 


X 


X 


9 


F granulose 










X 


X 




X 






3 


Pocillopora damicornis 






X 




X 


X 


X 


X 


X 


X 


8 


P. eydouxi 


X 


X 


X 


X 


X 


X 


X 


X 


X 




9 


P. sp.10 cf. laysanensis 






X 






X 








X 


4 


P. ligulata 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


11 


P. meandrina 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


10 


P. molokensis 


X 




X 


X 


X 


X 


X 


X 




X 


9 


P. sp.32 cf. verrucosa 






X 








X 


X 






3 


P. sp.33 cf. zelli 






X 
















1 


*P. sp.ll cf. capitata 






X 




X 


X 


X 


X 


X 


X 


8 


*Porites sp.12 cf. annae 














X 


X 




X 


3 


*P. sp. 15 (paliform lobes) 






X 
















1 


Pontes brighami 


X 


X 


X 


X 


X 


X 


X 


X 




X 


9 


P. compressa 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


11 


*P. sp.23 (arthritic fingers) 






X 
















1 


P. duerdeni 




X 


X 


X 


X 






X 




X 


6 


P. evermanni 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


10 


P. hawaiiensis 




X 


X 




X 


X 


X 


X 


X 


X 


8 


P. lobata 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


10 


*P. sp. 21 cf. lobata 






X 
















1 


*P. sp.16 cf. lutea 






X 
















1 


P. rus 










X 












1 


*P. sp.27 (columns) 






X 
















1 


*P. sp.13 cf. solida 






X 


X 




X 




X 


X 


X 


6 


Psammocora explanulata 








X 














1 


P. nierstraszi 




X 


X 


X 


X 


X 


X 


X 


X 




8 


P. stellata 


X 




X 


X 


X 


X 


X 


X 


X 


X 


10 


P. verrilli 
















X 


X 


X 


3 


Total 


17 


21 


66 


29 


41 


34 


37 


43 


33 


36 




Endemic 


4 


8 


27 


7 


12 


11 


15 


14 


8 


15 


Mean 


Percent endemic 


23.5 


38.1 


40.9 


24.1 


29.3 


32.3 


40.5 


32.6 


24.2 


41.7 


32.8 



Non-stony corals and anemones. 






















Non-Stony Corals & Anemones 


NIH 


MMM 


FFS 


GAR 


MAR 


LAY 


LIS 


PHR 


MID 


KUR 


Palythoa tuberculosa 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


P. sp. 






X 
















Zoanthus pacificus 






X 




X 






X 




X 


Zoanthus sp (Kure) 




















X 


Zoanthus sp ("B") 


X 


X 


X 




X 




X 








*Sinularia sp (yellow) 


X 


X 


X 


X 






X 








*Sinularia (purple) 








X 












X 


*Sinularia (brown) 






X 
















*Sinularia (pink) 
















X 






Acabaria bicolor 






X 














X 


Cirrhipathes sp 


X 




X 
















Heteractis malu 






X 


X 


X 


X 








X 


Total Species Per Island 


4 


3 


9 


4 


4 


2 


3 


3 


1 


6 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

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PERSONAL COMMUNICATIONS 

Maragos, J.E. U.S. Fish and Wildlife Service, Honolulu, HI, U.S.A. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Fishes 

Alan Friedlander 1 ' 2 , Edward DeMartini 3 , Lisa Wedding 14 and Randy Clark 1 



BIOGEOGRAPHY OF FISHES 

The Hawaiian Archipelago is among the most isolated on earth and exhibits the highest level of marine fish 
endemism of any archipelago in the Pacific (Randall, 1995, 1998, 2007; Randall and Earle, 2000; Allen, 2002). 
Owing to limited human influence, the Northwestern Hawaiian Islands (NWHI) reefs are nearly pristine and 
represent one of the last remaining intact large-scale, predator-dominated coral reef ecosystems on earth 
(Friedlander and DeMartini, 2002). Because of its high level of endemism and the near pristine nature of its 
reefs, the NWHI represents an important global biodiversity hot spot and provides a view of what reefs in the 
may have MHI looked like before human contact. 

Despite centuries of exploitation, the MHI today has even higher biodiversity of fishes than the NWHI. Randall 
et al. (1993) reported 258 species of reef and shore fishes from Midway Atoll compared with 612 species in 
the MHI (Randall, 2007). Mundy (2005) lists 21 species that are known from the NWHI, but not the MHI (Table 
5.1). Of these, most are either deep-water or mesopelagic and therefore poorly sampled waifs, or species with 
poor taxonomic resolution. In contrast, 406 species are known from the Main Hawaiian Islands (MHI) but not 
the NWHI and overall richness and diversity are greater in the MHI compared with the NWHI (Mundy, 2005). 



Table 5.1. Fish . 


species known from the NWHI but not found in the MHI. Source: Mundy, 2005. 




FAMILY 


SPECIES 


COMMON NAME 


LOCATIONS 


HABITAT 


Scyliorhinidae 


Apristurus spongiceps 


Spongehead catshark 


Nihoa 


Deep-water 


Muraenidae 


Gymnothorax atolli 


Atoll moray 


Pearl and Hermes to Midway 


Cryptic 


Platytroctidae 


Mentodus mesalirus 


Tubeshoulders 


Pearl and Hermes to Midway 


Deep-water 


Stomiidae 


Astronesthes nigroides 


Dragonfish 


Pearl and Hermes to Midway 


Mesopelagic 


Eustomias cancriensis 


Scaleless black dragonfish 


Pearl and Hermes to Midway 


Mesopelagic 


Ophidiidae 


Bassozetus zenkevitchi 


Cusk-eel 


Midway to Kure 


Deep-water 


Spectrunculus grandis 


Cusk-eel 


Maro 


Deep-water 


Macrouridae 


Cetonurus crassiceps 


Grenadier, Rattail 


Pearl and Hermes 


Deep-water 


Holocentridae 


Myripristis murdjan 


Blotcheye soldierfish 


Midway to Kure 


Shallow reefs 


Fistulariidae 


Fistularia petimba 


Serrate coronetfish 


Nihoa to Kure 


Mod-deep- 
water 


Laysan 


Scorpaenopsis pluralis 


Laysan scorpionfish 


Lasyan 


Deep-water 


Callanthiidae 


Grammatonotus macrophthalmus 


Splendid perch 


French Frigate Shoals 


Deep-water 


Epigonidae 


Epigonus devaneyi 


Deepwater cardinalfish 


Mokumanamanato Maro 


Deep-water 


Carangidae 


Caranx lugubris 


Black trevally 


Mokumanamanato Midway 


Shallow to 
deep 


Carangidae 


Decapterus macrosoma 


Shortfin scad 


Maro 


Pelagic 


Pomacanthidae 


Centropyge interruptra 


Japanese angelfish 


Kure and Midway 


Shallow reefs 


Kyphosidae 


Girella leonina 


Blackedge nibbler 


Midway 


Waif 


Labridae 


Epibulus insidiator 


Slingjaw wrasse 


French Frigate Shoals north 
to Kure 


Shallow reefs 


Ammodytidae 


Lepidammodytes macrophthalmus 


Sand lance 


Maro 


Poorly known 


Ephippidae 


Platax boersii 


Boer's spadefish 


Midway 


Waif 


Luvaridae 


Luvarus imperialis 


Louvar 


Laysan 


Epipelagic 



1. NOAA/NOS/NCCOS/CCMA Biogeography Branch 

2. The Oceanic Institute 

3. NOAA/NMFS/Pacific Islands Fisheries Science Center, Coral Reef Ecosystem Division 

4. University of Hawaii at Manoa 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Figure 5.1. The Japanese angelfish (Centropyge interruptra; left) and the 
spotted knifejaw (Oplegnathus punctatus; right) are known only from the 
NWHI and Japan, although the latter is occasionally observed in the MHI. 
Photo: J. Watt. 



The few demersal species found in the 
NWHI but not the MHI include the blotch- 
eye soldierfish (Myripristis murdjan), an 
Indo-Pacific species but restricted to the 
NWHI in the Hawaiian archipelago, and 
the Japanese angelfish (Centropyge 
interruptra; Figure 5.1) which is known 
only from the NWHI and Japan (Mundy, 
2005). Evidence of larval pelagic trans- 
port from Japan to the NWHI via the 
Kuroshio and North Pacific Currents is 
supported by the presence of a number 
of species that are common only off Ja- 
pan and the northwestern end of the Hawaiian Archipelago (Randall, 2007). In addition to the Japanese an- 
gelfish, these include two lizardfishes (Synodus lobelia and S. ulae), the manyspine squirrelfish (Sargocentron 
spinosissimum), two species of knifejaws (Oplegnathus fasciatus and O. punctatus) and the blackedge nibbler 
(Girella punctata , family Girellidae), a close relative of the chubs of the family Kyphosidae (Randall, 2007). 

Two species, the slingjaw wrasse (Epibulus insidiator) and the chevron butteryflyfish (Chaetodon trifascialis), 
are associated with Acropora corals that occur only in the central portion of the NWHI, and although these fish 
species are occasionally observed in the MHI and the far northern end of the chain, they are most abundance 
from French Frigate Shoals to Pearl and Hermes (Mundy, 2005). Despite the taxonomic similarity with the MHI 
fauna, the NWHI fish assemblage differs from that of the MHI at various ecological and demographic levels ow- 
ing to oceanographic conditions (e.g., water temperature), habitat (e.g., coral and reef type) and anthropogenic 
influences (e.g., effects of fishing in the MHI). 

There are a variety of environmental and 
other reasons for lower reef fish diversi- 
ty in the NWHI versus MHI. Many shal- 
low-water fish species that are adapted 
to warmer water cannot survive in the 
NWHI since winter water temperatures 
can be as much as 7°C cooler than the 
MHI (Mundy, 2005). Some shallow-wa- 
ter species are adapted to cooler water 
and can be found in deeper waters at 
the southern end of the archipelago. 
This phenomenon known as tropical 
submergence is exemplified by species 

such as the yellowfin soldierfish (Myripristis chryseres), the endemic Hawaiian grouper (Epinephelus quernus), 
and the masked angelfish (Genicanthus personatus), all of which occur in shallow water at Midway but are 
restricted to much greater depths in the MHI (Figure 5.2; Randall et al., 1993; Mundy, 2005). Other reasons for 
the lower number of species in the NWHI include insufficient sampling effort and the lack of many high island 
habitats such as estuaries and rocky shorelines 

Some of the non-endemic species abundant at higher latitude reefs in the NWHI have antitropical distribu- 
tions and are thought to have established themselves in the archipelago when surface waters were previously 
cooler (Randall, 1981). The Hawaiian morwong, (Goniistius vittatus) for example, may be a cryptic species that 
diverged during the late Miocene-early Pliocene from the lineage presently represented by nominal conspecif- 
ics in the southern hemisphere (Burridge and White, 2000). Interestingly, most Hawaiian endemic species do 
not appear to exhibit submergence (greater depth distributions) in the MHI, although rigorous comparisons are 
lacking (DeMartini and Friedlander, 2004). 




Figure 5.2. The endemic masked angelfish (Genicanthus personatus, left) 
and Hawaiian grouper (Epinephelus quernus, right) are found in shallower 
water at Midway Atoll but are restricted to deeper depths in the MHI. Pho- 
tos: J. Watt (left); J. Maragos (right). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Fish Species Richness 

Despite lower species richness in the 
NWHI as compared with the MHI (Mun- 
dy 2005), the total number of species 
(210) observed on quantitative transects 
in the NWHI (DeMartini and Friedlander, 
2004) was similar to the number of spe- 
cies (215) reported in a recent compre- 
hensive quantitative study around the 
MHI (Friedlander et al., 2007). The low- 
est overall fish species richness in the 
NWHI occurs at the small basalt islands 
(Mokumanamana, Gardner and Nihoa; 
Figure 5.3) and highest at French Frig- 
ate Shoals and Pearl and Hermes. The 
former may be related to the higher cor- 
al richness and greater diversity of habi- 
tats (Maragos et al., 2004), while the lat- 
ter is likely related to large size, habitat 
diversity and presence of subtropical 
and temperate species which occur at 
much greater depths southward in the 
chain of islands. 

Total species richness observed on 
surveys (y) showed a positive, linear 
relationship (y=8.05 * ln(x+l) + 112.2, 
R 2 =0.51, p=0.02, Table 5.2, Figure 5.4) 
with a logarithmic function of total reef 
area less than 10 fathoms (x). This re- 
lationship is consistent with the general 
theory of island biogeography and likely 
reflects the greater diversity of habitats 
present in larger reef areas. 




Figure 5.3. Total fish species richness at each of 10 emergent NWHI reefs. 
Source: NWHI RAMP, unpub. data; map: L. Wedding. 

Table 5.2. Results of least squares linear regression model for total num- 
ber of species by In (total reef area within 10 fathoms +1). R 2 = 0.51, N = 
10. Source: NWHI RAMP, unpub. data. 



ANALYSIS OF VARIANCE 


Source 


DF 


Mean Square 


F Ratio 


Prob > F 


Model 


1 


2105.32 


8.20 


0.0211 


Error 


8 


256.85 






C. Total 


9 








PARAMETER ESTIMATES 


Term 


Estimate 


Standard Error 


t Ratio 


Prob>|t| 


Intercept 


112.21 


12.3 


9.12 


<0.001 


Ln (area) 


8.052 


2.81 


2.86 


0.0211 



CD 

o 

CD 
Q. 
If) 

M — 

o 

CD 
-Q 

E 

"tC 

o 

H 


220 - 
200 - 
180 - 
160 - 
140 - 
120 - 
100 - 
80 - 
60 - 

40 - 

t 








• ^ — ——" 




__ — - — • _^^- — 




• Observations 

Linear regression 

95% Confidence Band 

95% Prediction Band 


) 2 4 6 

ln[(reef area (km 2 ) < 10 fm) + 1] 





Figure 5.4. Relationship between cumulative number of fish species at 
each reef and total reef area (km 2 ) within 10 fathoms. Source: Friedlander 
et al., in prep. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Endemism 

The Hawaiian Island chain is among the most isolated on earth and exhibits the highest level of marine fish 
endemism of any archipelago in the Pacific (Randall, 1995, 1998; Randall and Earle, 2000; Allen, 2002). En- 
demism is a key attribute of biotic communities that is generally a great concern of conservation ecology. One 
reason of general biogeographic interest is that speciation and the origin and maintenance of biodiversity are 
undoubtedly related to degrees of isolation and endemism (Gray, 1997). Because of the decline in global ma- 
rine biodiversity, endemic "hot spots" like Hawaii are important areas for global biodiversity conservation. The 
endemic fishes of Hawaii are small bodied and have very restricted geographic ranges of less than 50,000 km 2 
(Roberts et al., 2002). Small body size, per se, may be associated with higher extinction risk because small- 
bodied species tend to have narrower habitat requirements (Hawkins et al., 2000). Therefore both body size 
and endemic status argue for the conservation of these species. 

Based on species-presence, endemism is equivalent for fishes in the NWHI (20.6% using all available data) 
and the MHI (MHI, 20.9%; DeMartini and Friedlander, 2004). On average, percentage endemism was much 
higher based on numerical densities (52%) and biomass (37%) which increased with latitude, and was espe- 
cially pronounced at the four northernmost reefs that are the oldest emergent geological features of the archi- 
pelago (Figure 5.5). Greater endemism towards Midway and Kure appears related to consistently higher rates 
of replenishment by young-of-the-year (YOY) upchain following dispersal as pelagic larvae and/or juveniles. 
There were significant positive relationships between number and biomass of endemics with latitude (Tables 
5.3, 5.4, 5.5; Figures 5.6). However endemism based on species presence was not significantly correlated with 
latitude (Figure 5.7). 





Figure 5.5. Percent endemism based on numerical densities (top left), biomass (top right) and species richness (bottom 
left) at each of 10 emergent NWHI reefs. Source: DeMartini and Friedlander, 2004; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Endemic reef fishes are appreciably smaller bodied than nonendemics within the NWHI (DeMartini and Fried- 
lander, 2004). Median body size does not vary with latitude and longitude for either endemics or nonendem- 
ics, which obviates possibly confounding environmental effects. Reef fish populations at higher latitude reefs 
included larger proportions of YOY recruits. YOY length frequencies did not differ for most species between 
northern and southern reefs, suggesting that a seasonal lag in spawning and recruitment at higher latitudes 
cannot explain the greater YOY densities observed. Disproportionate recruitment at higher-latitude reefs may 
be related to better growth and survivorship after settlement onto reefs, higher levels of within-reef and re- 
gional reseeding at higher latitudes, or other factors. 



Table 5.3. Results of least squares linear regression model 
for total number of species by In (total reef area within 10 
fathoms +1). R 2 = 0.51, N = 10. Source: DeMartini and 
Friedlander, 2004. 



ANALYSIS OF VARIANCE 


Source 


DF 


Mean 
Square 


F Ratio 


Prob > F 


Model 


1 


962.8 


15.86 


0.004 


Error 


8 


60.72 






C. Total 


9 









PARAMETER ESTIMATES 
Estimate 



Term 

Intercept 



Latitude 



-89.86 



5.28 



Standard 
Error 

34.15 



1.32 



t Ratio Prob>|t| 

-2.63 0.0301 



3.98 



0.004 



Table 5.5. Results of least squares linear regression model 
for total number of species by In (total reef area within 10 
fathoms +1). R 2 = 0.51, N = 10. Source: DeMartini and 
Friedlander, 2004. 



ANALYSIS OF VARIANCE 


Source 


DF 


Mean 
Square 


F Ratio 


Prob > F 


Model 


1 


8.18 


3.53 


0.0971 


Error 


8 


2.32 






C. Total 


9 








PARAMETER ESTIMATES 


Term 


Estimate 


Standard 
Error 


t Ratio 


Prob>|t| 


Intercept 


13.77 


6.67 


2.06 


0.0729 


Latitude 


0.49 


0.26 


1.88 


0.0971 



Table 5.4. Results of least squares linear regression model 
for total number of species by In (total reef area within 10 
fathoms +1). R 2 = 0.51, N = 10. Source: DeMartini and 
Friedlander, 2004. 



ANALYSIS OF VARIANCE 


Source 


DF 


Mean 
Square 


F Ratio 


Prob > F 


Model 


1 


1817.66 


14.7374 


0.005 


Error 


8 


123.34 






C. Total 


9 








PARAMETER ESTIMATES 


Term 


Estimate 


Standard 
Error 


t Ratio 


Prob>|t| 


Intercept 


-153.734 


48.67219 


-3.16 


0.0134 


Latitude 


7.249942 


1.888528 


3.84 


0.005 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 





100 - 












• Observations 








Linear regression line 






3 
T3 




95% Confidence Band 






> 


80 - 


95% Prediction Band 






T3 
E 










o 










CD 
.Q 
E 

3 


60 - 


• ___ — — — ' 


• ^^~~-^ 




E 

CO 


40 - 








E 

CD 

E 
LLI 




•^^^^ ^-~~* 






E 
CD 

CJ 

CD 
CL 


20 - 

- 


^*^m ^^~^^ 














2 


2 24 


26 
Latitude 


28 





100 -i 








• Observations 








Linear regression line 




en 


80 - 


95% Confidence Band 




ns 
E 

o 




95% Prediction Band ^^_^- 




60 - 






CO. 




^____— -■—-■■■"'" • 


-^"^ • — -"-■"■-"""""""""^ 










u 








F 


40 - 






en 








E 




— — — — -~~~ — ~~ — *~ ^^^^^\^-^^" — ""~~~~ 




CD 

•o 


20 - 


— - -— """"""'"''"'^ ^*^* - 




LU 
















E 
CD 


- 






O 
















CD 








0_ 


-20 - 
-40 - 












22 24 26 


28 




Latitude 





Figure 5.6. Least squares linear regression model for percent endemism by numerical abundance versus latitude (left). 
Least squares linear regression model for percent endemism by biomass versus latitude (right). Source: DeMartini and 
Friedlander, 2004. 



'o 

CD 
Q. 
CO 

E 
S 

<D 

T3 
C 
LU 



0) 

o 



ID 
CL 





• 


Observations 












29 - 
















28 - 






• 








• 


27 - 




• 






• 






26 - 




• 




• 


• 


• 




25 - 
















24 - 
















23 - 




• 












?? - 




, , 


. 




. 


. 





22 



23 



24 



25 26 

Latitude 



27 



28 



29 



Figure 5. 7. Least squares linear regression model for percent endemism by 
species versus latitude. Source: DeMartini and Friedlander, 2004 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



LATITUDINAL AFFINITIES AMONG FISHES 



Table 5.6. Percentage of numerical abundance at each reef that consisted 
of species that showed either a temperate/subtropical (northerly) affinity 
or tropical (southerly) affinity in abundance. Source: Friedlander et al., in 
prep. 



REEF 


TEMPERATE/SUBTROPICAL AFFINITY 


TROPICAL AFFINITY 


NIH 


12.97% 


16.35% 


MMM 


28.01% 


28.44% 


FFS 


27.45% 


8.57% 


GAR 


24.52% 


14.15% 


MAR 


51.94% 


4.27% 


LAY 


44.91% 


10.24% 


LIS 


52.90% 


3.22% 


PHR 


52.34% 


1.99% 


MID 


56.08% 


0.93% 


KUR 


63.43% 


1.24% 


Island/atoll abbreviations used throughout this chapter: NIH = Nihoa Island; MMM = 
Mokumanamana, FFS = French Frigate Shoals; GAR = Gardner Pinnacles; MAR = Maro 
Reef; LAY = Laysan Island; LIS = Lisianski Island; PHR = Pearl and Hermes; MID = Mid- 
way Atoll; KUR = Kure Atoll 



Biogeographic forces may promote dis- 
parate abundance patterns among some 
species at opposite ends of the archipel- 
ago owing to differences in temperature 
and other environmental factors. Some 
species might have a temperate or sub- 
tropical bias, whereas others might be 
better suited to more tropical conditions. 
To identify latitudinal gradients of abun- 
dance, numerical densities as a func- 
tion of latitude was examined within the 
NWHI using Spearman rank correlation. 
Positive correlations indicated a temper- 
ate affinity, while negative correlations 
indicated a tropical affinity. The percent- 
age of individuals with either temperate/ 
subtropical or temperate affinities is an 
indication of the total fish assemblage 
affinity at each reef (Table 5.6; Figure 
5.8). 

Thirty species showed a significant pos- 
itive correlation (Spearman Rank Cor- 
relation, p<0.05) with latitude based on 
numerical density from quantitative fish 
surveys conducted between 2000 and 
2002 (Table 5.7). Of these, 17 (57%) 
were endemics. Wrasses (Labridae) had 
the greatest number of species (eight) 
showing higher latitude affinity followed 
by damselfishes (Pomacentridae) with 
four species. Several other species 
such as knifejaws (Oplegnathus spp.) 
and boarfish (Evistias acutirostris) were 
more abundant at higher latitudes but 
their low numbers during surveys made 
the results inconclusive statistically. 

1T5TW 17im 16j*W 

Figure 5.8. Percentage of total numerical abundance (numbers m 2 ) for spe- 
cies showing a significant latitude gradient of distribution. Source: Fried- 
lander et al., in prep. 

Table 5.7. Species with temperate/subtropical affinity (positive correlation with latitude). Endemics in bold. Source: 
Friedlander et al., in prep. 




FAMILY 


TAXON NAME 


COMMON NAME 


HAWAIIAN NAME 


Synodontidae 


Synodus ulae 


Ulae Lizardfish 


ulae 


Holocentridae 


Sargocentron xantherythrum 


Hawaiian Squirrelfish 


alaihi 


Scorpaenidae 


Pterois sphex 


Hawaiian Turkeyfish 




Serranidae 


Epinephelus quernus 


Hawaiian Grouper 


hapuu 


Priacanthidae 


Priacanthus meeki 


Hawaiian Bigeye 


aweoweo 


Chaetodontidae 


Chaetodon auriga 


Threadfin Butterflyfish 


kikakapu 


Pomacanthidae 


Genicanthus personatus 


Masked Angelfish 




Pomacentridae 


Abudefduf abdominalis 


Sargent Major 


mamo 


Pomacentridae 


Chromis hanui 


Chocolate-dip Chromis 






A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 5.7 (continued). Species with temperate/subtropical affinity (positive correlation with latitude). Endemics in bold. 
Source: Friedlander et al., in prep. 


FAMILY 


TAXON NAME 


COMMON NAME 


HAWAIIAN NAME 


Pomacentridae 


Chromis ovalis 


Oval Chromis 




Pomacentridae 


Stegastes fasciolatus 


Pacific Gregory 




Cirrhitidae 


Paracirrhites forsteri 


Blackside Hawkfish 


hilu pili koa 


Labridae 


Anampses cuvier 


Pearl Wrasse 


opule 


Labridae 


Coris flavovittata 


Yellowstrip coris 


hilu 


Labridae 


Gomphosus varius 


Bird Wrasse 


hinaleaiiwi, akilolo 


Labridae 


Labroides phthirophagus 


Hawaiian Cleaner Wrasse 




Labridae 


Stethojulis balteata 


Belted Wrasse 


omaka 


Labridae 


Thalassoma ballieui 


Blacktail Wrasse 




Labridae 


Thalassoma duperrey 


Saddle Wrasse 


hinalea lauwili 


Labridae 


Thalassoma purpureum 


Surge Wrasse 


hou 


Scaridae 


Calotomus zonarchus 


Yellowbar Parrotfish 




Scaridae 


Chlorurus perspicillatus 


Spectacled Parrotfish 


uhu uliuli 


Scaridae 


Scarus dubius 


Regal Parrotfish 


lauia 


Cheilodactylidae 


Cheilodactylus vittatus 


Hawaiian Morwong 




Acanthuridae 


Acanthurus nigroris 


Bluelined Surgeonfish 


maiko 


Acanthuridae 


Zebrasoma veliferum 


Sailfin tang 


maneoneo 


Gobiidae 


Coryphopterus sp. 


Goby 


oopu 


Gobiidae 


Cnatholepis anjerensis 


Eyebar goby 




Balistidae 


Xanthichthys mento 


Crosshatch Triggerfish 




Diodontidae 


Diodon holocanthus 


Spiny Puffer 


oopu okala 



Over 63% of the total numerical abundance of fishes at Kure Atoll was composed of species with a high lati- 
tude correlation (Figure 5.8). The percentage of high latitude affinity individuals was also substantial at Midway 
Atoll (56%), Pearl and Hermes Atoll (52%) and Lisianski Island-Neva Shoals (53%). The major break occurs 
between Maro Reef and Gardner Pinnacle where the numerical abundance of high latitude affinity species 
dropping from 52% to 25% between these two locations. The lowest percentage of high latitude affinity indi- 
viduals was observed at Nihoa Island (13%). There was a relatively large shift towards more high latitude af- 
finity individuals between Nihoa and Mokumanamana (28%). 

Twenty-one species were significantly and positively correlated (p<0.05) with low latitudes based on numerical 
density estimated on surveys conducted between 2000-2002 (Table 5.8). Only two of these species (9%) were 
endemics in contrast to the species with high latitude bias, where 54% were found to be endemic. Based on 
total numerical abundance, the highest percentage of low latitude species was observed at Mokumanamana 



Table 5.8 Species with tropical affinity (negative correlation with latitude). Endemics in bold. Source: Friedlander et al., 
in prep. 



FAMILY 


TAXON NAME 


COMMON NAME 


HAWAIIAN NAME 


Carcharhinidae 


Carcharhlnus amblyrhynchos 


Gray Reef Shark 


mano 


Carcharhinidae 


Triaenodon obesus 


Whitetip Reef Shark 


mano lalakea 


Lutjanidae 


Aphareus furca 


Smalltooth Jobfish 


wahanui 


Lethrinidae 


Monotaxis grandoculis 


Bigeye Emperor 


mu 


Mullidae 


Parupeneus bifasclatus 


Doublebar Goatfish 


munu 


Mullidae 


Parupeneus multifasciatus 


Manybar Goatfish 


moano 


Chaetodontidae 


Chaetodon multicinctus 


Multiband Butterflyfish 


kikakapu 


Chaetodontidae 


Chaetodon quadrimaculatus 


Fourspot Butterflyfish 


lau hau 


Pomacentridae 


Plectroglyphidodon imparipennis 


Brighteye Damselfish 




Cirrhitidae 


Paracirrhites arcatus 


Arc-eye Hawkfish 


pili koa 


Scaridae 


Calotomus carolinus 


Stareye Parrotfish 




Acanthuridae 


Acanthurus blochii 


Ringtail Surgeonfish 


pualu 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Table 5.8 (continued). Species with tropical affinity (negative correlation with latitude). Endemics in bold. Source: Fried- 
lander et al., in prep. 



FAMILY 


TAXON NAME 


COMMON NAME 


HAWAIIAN NAME 


Acanthuridae 


Acanthurus nigrofuscus 


Brown Surgeonfish 


maiii 


Acanthuridae 


Acanthurus olivaceus 


Orangeband Surgeonfish 


naenae 


Acanthuridae 


Naso lituratus 


Orangespine Unicornfish 


umaumalei 


Monacanthidae 


Cantherhines sandwichiensis 


Squaretail Filefish 


oili lepa 


Balistidae 


Melichthys niger 


Black Durgon 


humuhumuelele 


Monacanthidae 


Pervagor aspricaudus 


Lacefin Filefish 




Balistidae 


Rhinecanthus rectangulus 


Reef Triggerfish 


humuhumunukunukuapuaa 


Balistidae 


Sufflamen bursa 


Lei Triggerfish 


humuhumulei 


Tetraodontidae 


Canthigaster amboinensis 


Ambon Toby 





(28%) and Nihoa (14%; Figure 5.8). Less 
than 1% of the number density of fishes 
counted at Midway consisted of species 
with a low latitude preference. Similar- 
ly, Kure Atoll (1.2%) Pearl and Hermes 
Atoll (2.0%) and Lisianski Island-Neva 
Shoals (3.2%) had low numbers of more 
tropical affinity individuals. 

There is a strong positive linear rela- 
tionship between the percentage of 
individuals with temperate/subtropical 
affinities and latitude (Table 5.9, Figure 
5.9), while there is a strong negative 
linear relationship with the percentage 
of individuals with tropical affinities and 
latitude (Table 5.10, Figure 5.9). A ma- 
jor faunal break occurred around Maro 
and Laysan, where the numerical abun- 
dance of northern and southern affin- 
ity species were more similar. Although 
species with northern affinities were still 
more abundant than species with south- 
ern affinities south of Maro, the overall 
numerical abundance of these northern 
species averaged 23% south of Maro, 
but 54% to the north. Species with tropi- 
cal affinities account for 17% of fish 
numbers south of Maro, but only 4% to 
the north. 



Table 5.9. Least squares linear regression model for species exhibiting tem- 
perate/subtropical affinity and latitude. Source: Friedlander et al., in prep. 



ANALYSIS OF VARIANCE 


Source 


DF 


Mean 
Square 


F Ratio 


Prob > F 


Model 


1 


0.20 


33.57 


0.0004 


Error 


8 


0.01 






C. Total 


9 








PARAMETER ESTIMATES 


Term 


Estimate 


Standard 
Error 


t Ratio 


Prot»|t| 


Intercept 


-1.57 


0.34 


-4.57 


0.0018 


Latitude 


0.08 


0.01 


5.79 


0.0004 





1.0 -| 








# Temperate/subtropical 








# Tropical 








^^— Linear regression temperate 






0.8 - 


95% Confidence Band 




o 




^^— Linear regression tropical 




a 




95% Confidence Band 




E 
3 


0.6 - 






.Q 






^*"*^*» — - 


< 




• 5-<^ 


* — 1^-^"~"~^ 


IIS 








o 




^ — ^-~- 




ID 


0.4 - 






E 








3 








Z 




^-^cr*^-- — *^^ 




C 




~^^Z^^ m 




t 


0.2 - 















o. 

























a. 


0.0 - 








1 ' 


1 




22 24 26 


28 




Latitude 





Figure 5.9. Relationship between latitude and numerical abundance of 
species with temperate/subtropical and tropical affinities. Results of least 
squares linear regression. Temperate/subtropical = -1.56 + 0.07*Latitude, 
Tropical = 1.00 - 0.03*Latitude. Source: Friedlander et al., in prep. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Fish Recruitment 

The planktonic dispersal of reef fishes 
is an important process linked to the 
persistence of benthic reef populations. 
Recruitment of reef fishes increased 
with latitude, and was especially pro- 
nounced at the four northernmost reefs 
that had a larger proportion of YOY re- 
cruits (DeMartini and Friedlander, 2004). 
During 2000-2002, recruit fish densities 
were generally greater upchain to the 
northwest (versus downchain) and a 
larger number of endemic (versus non- 
endemic) species recruited to a great- 
er extent upchain in the NWHI (Figure 
5.10; DeMartini and Friedlander, 2004). 
YOY recruit length frequencies did not 
differ for most species between north- 
ern and southern reefs, suggesting that 
a seasonal lag in spawning and recruit- 
ment at higher latitudes cannot explain 
the greater YOY densities observed 
there. Disproportionate recruitment at 
higher-latitude reefs may be related to 
higher levels of within-reef and region- 
al reseeding at higher latitudes. This 
was first indicated by survey data col- 
lected during the 1990s at French Frig- 
ate Shoals and Midway (DeMartini et 
al., 2002; DeMartini, 2004). During this 
period, there was consistently higher 
recruitment of YOY life stages of fishes 
at Midway Atoll versus French Frigate 
Shoals despite the generally greater 
densities of older-stage fishes at French 
Frigate Shoals (Figure 5.11). 



Table 5.10. Least squares linear regression model for species with tropical 
affinity and latitude. Source: NWHI RAMP, unpub. data. 



ANALYSIS OF VARIANCE 


Source 


DF 


Mean 
Square 


F Ratio 


Prob > F 


Model 


1 


0.04 


14.17 


0.0055 


Error 


8 


0.01 






C. Total 


9 








PARAMETER ESTIMATES 


Term 


Esti- 
mate 


Standard 
Error 


t Ratio 


Prob>|t| 


Intercept 


1.01 


0.24 


4.12 


0.0033 


Latitude 


-0.04 


0.01 


-3.76 


0.0055 




Figure 5. 1 0. Geographic patterns of the Recruit Index (ratio of YO Y sized to 
larger individuals) for all pooled major species of endemic and non-endemic 
reef fishes. Source: DeMartini and Friedlander, 2004; map: L. Wedding. 




1992 1993 1994 1995 1996 1997 1998 1999 2000 



Figure 5.11. Time series of the estimated mean numerical density of YOY 
of all taxa at French Frigate Shoals and Midway during each survey year. 
Each vertical bar represents one southeast of the estimated survey year 
grand mean for both major habitats. Source: DeMartini, 2004. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



GENERAL FISH ASSEMBLAGE STRUCTURE 

Dominance by species was revealed by 
plotting relative percent contribution by 
each species to total biomass at each 
reef. A limited number of species ac- 
counted for the majority of the biomass 
for most locations. Giant trevally (ulua, 
Caranx ignobilis) was the dominant spe- 
cies by weight at Lisianski (50% of to- 
tal biomass), Pearl and Hermes (43%), 
Laysan (32%) and Maro (30%; Figure 
5.12). Chub (nenue, Kyphosus spp.) 
is the most dominant taxa by weight at 
Nihoa and accounts for 35% of the bio- 
mass. 



The similarity of fish assemblages 
among reefs in the NWHI was com- 
pared based on biomass density for 
each species at each reef (Figures 5.13, 
5.14). Two atolls (Kure and Midway) had 
high concordance and formed a distinct 
cluster relative to all other islands. The 
two basalt islands (Nihoa and Mokuma- 
namana) were also distinct in their fish 
assemblages while Gardner Pinnacles, 
the other basalt rock, was unique in its 
fish assemblage based on biomass. 
Pearl and Hermes and Lisianski were 
the most similar based on fish assem- 
blage biomass but also cluster at lower 
levels with Maro, Laysan, and to a less- 
er extent, French Frigate Shoals. 

Similarity based on numerical abun- 
dance showed two distinct clusters with 
Nihoa being an extreme outlier (Fig- 
ures 5.15, 5.16). Midway and Pearl and 
Hermes exhibited similar assemblage 
structure, as did French Frigate Shoals 
with Maro, and Kure with Lisianski. Mo- 
kumanamana, Gardner, and to a lesser 
extent, Laysan, exhibited similar assem- 
blage structure but were less correlated 
than those in the other cluster. Nihoa 
was unique in its assemblage structure 
based on numerical abundance. 




Nihoa 

Mokumanamana 

French Frigate Shoals 

Gardner 

Maro 

Laysan 

Lisianski 

Pearl and Hermes 

Midway 

Kure 



10 
Species Rank 



Figure 5.12. Ordinary dominance curve for each reef based on biomass. 
Source: NWHI RAMP, unpub. data. 



20 -r 



E 



nc-- 



Transform: Log(X+l) 

Resemblance: S17 Bray Curtis Similarity 



Figure 5.13. Bray Curtis similarity dendrogram showing similarities among 
reef based on biomass. Source: NWHI RAMP, unpub. data. 



Stress: 0.07 




Figure 5.14. Nonmetric multi-dimensional scaling plot of reef similarities 
derived from biomass abundance of species. Similarities based on Bray- 
Curtis Similarity Index. Biomass abundance ln(x+l) transformed. Source: 
NWHI RAMP, unpub. data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



70 1 
75- 

80- 

>> 
'^ 

cti 

= 85- 

E 
in 

90- 
95- 

100- 






Transform: Fourth Root 

Resemblance: S17 Bray Curtis Similarity 


" 












1 










1 














I I 










1 






: Q rt 


1 

OT CC IE » > 
IL 3 Z) J « 

u- 5 * 



Figure 5.15. Bray Curtis similarity dendrogram showing similarities among 
reef based on numerical abundance. Source: NWHI RAMP, unpub. data. 



Trophic Structure 

Overall, apex predators accounted for 
47% of total fish biomass, followed by 
herbivores (31%) and secondary con- 
sumers (22%). Pearl and Hermes had 
the highest percentage of apex preda- 
tors (67%), with French Frigate Shoals 
(61%) and Lisianski-Neva Shoal (58%) 
also having substantial apex predator 
biomass (Figure 5.17). More than 65% 
of the apex predator biomass observed 
within the NWHI consisted of giant tre- 
vally. 

Apex predator biomass increases up the 
chain reaching a maximum at Pearl and 
Hermes Atoll before declining dramati- 
cally at Midway and Kure atolls (Figure 
5.17; DeMartini and Friedlander, 2004). 
The extremely low biomass of apex 
predators at Midway and Kure has been 
attributed to previous extractive fishing 
activities at both locations as well as a 
tag-and-release recreational sport fish- 
ery at Midway (DeMartini et al., 2002; 
DeMartini etal., 2005). 

Herbivores were dominant in terms of 
biomass at Nihoa (56%) and Midway 
(56%). Chubs accounted for most of the 
herbivore biomass at Nihoa while the 
endemic spectacled parrotfish (Chloru- 
rus perspicillatus) was most predomi- 
nant at Midway and Kure. The lowest 
total biomass was recorded at Moku- 
manamana (0.45 t ha 1 ) while Pearl and 
Hermes had the lowest percentage of 
herbivores (11%). Secondary consumer 

biomass ranged from a high at Midway (0.91 1 ha _1 ) and Pearl and Hermes (0.89 t ha - 1 ) to a low at Mokumana- 
mana (0.26 1 ha 1 ). The saddle wrasse was the dominant species among secondary consumers at both Midway 
and Pearl and Hermes. 




Figure 5.16. Nonmetric multi-dimensional scaling plot of reef similarities 
derived from numerical abundance of species. Similarities based on Bray- 
Curtis Similarity Index. Numerical abundance fourth root transformed. 
Source: NWHI RAMP, unpub. data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 






Figure 5.17. Percent biomass by consumer groups at each reef. Bubbles are proportional to total biomass (t ha 1 ). 
Source: NWHI RAMP, unpub. data. 



Biomass Size Spectra 
Biomass densities of pooled taxa were 
evaluated as size spectra relative to 
standardized length classes; our analy- 
sis revealed that there were relatively 
more and greater numbers of large in- 
dividual fish at Pearl and Hermes and 
French Frigate Shoals than elsewhere 
in the NWHI (Figure 5.18). Overall, 
biomass based on the intercept of the 
biomass-to-body size relation (i.e. 
abundance at the midpoint of the length 
distribution) was lowest at Kure, Moku- 
manamana and Nihoa (Figure 5.19). 





3.0 - 








Kure 










Midway 

PHR 




^^ 


2.5 - 




Lisianski 




'cfl 






Laysan 

Maro 






2.0 - 


^5^^^ 


Gardner 

FFS 




J^ 






Mokumanamana 




C/) 

E 
o 


1.5 - 
1.0 - 




Nihoa 








n 








o 

o 

_l 


0.5 - 
0.0 - 








-1 


DO -50 


50 100 






Standardized size class 


; (cm) 



Figure 5.18. Biomass size spectra for all fishes greater than 15 cm at each 
of the major reefs in the NWHI. Source: NWHI RAMP, unpub. data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Shark Distribution Patterns 
Sharks are an important component of 
the reef fish assemblage in the NWHI 
accounting for 28% of apex predator 
biomass and 13% of total reef fish bio- 
mass on the fore reef. Grey reef (Car- 
charhinus amblyrhynchos, 8.4%), Gala- 
pagos (Carcharhinus galapagensis, 
10.2%; Figure 5.20) and whitetip reef 
sharks (Triaenodon obesus, 8.6%; Fig- 
ure 5.20) comprised similar percentag- 
es of total apex predator biomass while 
blacktip reef sharks (Carcharhinus mel- 
anopterus) were much less abundant 
than the other three species and com- 
prised only 0.3% of total biomass and 
0.6% of apex predator biomass in the 
fore reef habitats. The biogeographic 
distribution patterns of gray reefs and 
Galapagos sharks were markedly differ- 
ent within the NWHI (Figure 5.21). Gray 
reef sharks were replaced by Galapa- 
gos sharks moving northward along the 
NWHI chain. Galapagos sharks are less 

abundant at Nihoa, Mokumanamana, and French Frigate Shoals but are very abundant northwest of Gardner 
Pinnacles. Gray reefs become less abundant northward. Papastamatiou et al. (2006) examined data from 
the Hawaii Shark Control Program between 1967 and 1980 and found Galapagos and tiger sharks (Galeor- 
cerdo cuvier) to be more abundant in the NWHI compared to the MHI, while sandbar sharks (Carcharhinus 
plumbeus) were more common in the MHI compared with the NWHI. These data showed gray reef sharks 
were more numerous in the NWHI compared with the MHI. Within the NWHI, this species was more abundant 
at Mokumanamana and French Frigate Shoals at the lower end of the NWHI and less abundant at the northern 
reefs of Maro and Midway. Interspecific competition, owing to dietary overlap, perhaps influences the distribu- 
tion of these sharks throughout the Hawaiian Islands (Papastamatiou et al., 2006). 




^vv.^ 



o.oio e» 

0.012 £& 



Figure 5.19. Plot of slope and y-intercept from size spectra regressions. 
Source: NWHI RAMP, unpub. data. 




Figure 5.20. Galapagos sharks (left) and a whitetip shark (right). Photos: J. Maragos and A. Friedlander. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 







Figure 5.21. Biogeographic distribution of sharks in the NWHI based on number of individuals ha 1 . Values are numbers 
ha 1 and are for fore reef habitats only. Source: NWHI RAMP, unpub. data. 



Influence of Influence of Predators 
on Prey Fishes 

The effects of apex predation, primar- 
ily by giant trevally, are pervasive. Apex 
predators structure prey population siz- 
es and age distributions and strongly in- 
fluence the reproductive and growth dy- 
namics of other harvested species (such 
as parrotfish) as well as smaller-bodied, 
lower-trophic-level fishes on shallow 
NWHI reefs (DeMartini and Friedlander, 
2006). Perhaps the strongest evidence 
for the controlling influence of apex pre- 
dation on the structure of fish assem- 
blages in the NWHI is provided by data 
on the size, composition and spatial dis- 
tribution of prey species (Figure 5.22; 
DeMartini etal., 2005). 





6 


- ■ • 

KUR 




J3 


5 


■ - i 




1 




MAR 




c 
g 

o 
D. 


4 
3 


- • ■ 
MID 

• FFS 

■ ■ 


PHR 


c 

or 


2 




▲ 


A 

• 


A 




# Median TL at sex change 


LIS 




1 




■ Median TL of all select labroids 
A Median TL of all other prey fishes 




1 










12 3 4 5 


6 




Rank Giant Trevally Density 





Figure 5.22. Scatterplot of the ranks of prey population attributes (median 
body length at sex change in the four major labroid species, median body 
lengths of all eight select species of labroids, and median body length of 
all other prey fishes) versus the ranks of giant trevally densities. Source: 
Demartinietal., 2005. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



DeMartini (2004) documented the 
habitat-specific spatial distributions of 
juvenile and other small-bodied fishes 
particularly susceptible to predation and 
recognized the importance of back reef, 
lagoon patch reef and other sheltered 
(wave-protected) habitats as nursery ar- 
eas for juvenile reef fishes (Figure 5.23). 
This study, based on re-analyses of data 
collected at French Frigate Shoals and 
Midway Atoll during the 1990s, has con- 
tributed substantially to development of 
both "essential fish habitat" and "habitat 
areas of particular concern" concepts 
in recognizing the greater per-unit-area 
value of atolls due to their larger propor- 
tion of sheltered juvenile nursery habi- 
tats (DeMartini, 2004). 




Figure 5.23. Percentage contribution of YOY to overall YOY plus older- 
stage densities. Source: adapted from DeMartini, 2004. 



Table 5.11. Fish species richness Analysis of Variance among 
Source: NWHI RAMP, unpub. data. 


r eefe. 


SOURCE 


DF 


SUM OF 
SQUARES 


MEAN 
SQUARE 


F RATIO 


PROB > F 


Reef 


9 


686.699 


76.3 


1.64 


0.1005 


Error 


400 


18548.44 


46.37 






C. Total 


409 


19235.14 









Updated Comparison of Fish Assemblage Metrics Among NWHI Reefs 
Based on data collected from initial sur- 
veys in 2000, 2001, 2002 and new sites 
surveyed in 2007, fish assemblage char- 
acteristics were compared among all 
reefs. Fish species richness appeared 
highest at Nihoa, Gardner and Laysan 
and lowest at Mokumanamana, Maro 
and Kure but these differences were not 
significant (Table 5.11, Figure 5.24). The 
number of individual fishes observed on 
transects differed significantly different 
among reefs (Table 5.12, Figure 5.25,). 
Midway, followed by Pearl and Hermes 
had the highest number of individuals 
while Mokumanamana and Maro had 
the lowest. Biomass also differed signifi- 
cantly different among reefs (Table 5.13, 
Figure 5.26; F g40g = 3.64, p < 0.001) with 
the highest biomass at Gardner, Nihoa 
and Pearl and Hermes. The lowest fish 
biomass was recorded at Kure and Mo- 
kumanamana. 




Figure 5.24. Mean species richness per transect from RE A data from 2000- 
2002 and 2007. Error bars are standard error of the mean. Source: NWHI 
RAMP, unpub. data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 5.12. Fish biomass in tha-1 (ln[x+l]) Analysis of 
Variance among reefs. Comparisons for all pairs using 
Tukey-Kramer HSD. Levels not connected by same letter 
are significantly different. Source: NWHI RAMP. 



Table 5.13. Fish biomass in tha-1 (ln[x+l]) Analysis of 
Variance among reefs. Comparisons for all pairs using 
Tukey-Kramer HSD. Levels not connected by same letter 
are significantly different. Source: NWHI RAMP. 



ANALYSIS OF VARIANCE 


Source 


DF 


Sum of 
Squares 


Mean 
Square 


F 
Ratio 


Prob 
>F 


Reef 


9 


6.78 


0.75 


6.18 


•c.OOOl 


Error 


400 


48.80 


0.12 






C. Total 


409 


55.58 








Level 


Multiple 
Comparisons 


Mean 








Midway 


A 


1.20 








PHR 


AB 


0.99 








Lisianski 


BC 


0.87 








FFS 


BC 


0.86 








Gardner 


ABC 


0.83 








Kure 


BC 


0.83 








Laysan 


BC 


0.83 








Nihoa 


BC 


0.81 








Maro 


BC 


0.79 








MMM 


C 


0.58 









ANALYSIS OF VARIANCE 


Source 


DF 


Sum of 
Squares 


Mean 
Square 


F 
Ratio 


Prob 
>F 


Reef 


9 


9.30 


1.03 


3.64 


0.0002 


Error 


400 


113.64 


0.28 






C. Total 


409 


122.93 








Level 


Multiple Com- 
parisons 


Mean 








Gardner 


AB 


1.26 








Lisianski 


A 


1.14 








Laysan 


AB 


1.13 








Midway 


A 


1.12 








Nihoa 


AB 


1.11 








PHR 


A 


1.09 








FFSs 


A 


1.01 








Maro 


AB 


0.97 








MMM 


AB 


0.74 








Kure 


B 


0.70 









1.4 -I 




F 9409 = 6.18; p < 0.001 


1.2 - 


T 


















r-i 
















& 1.0 - 






^ 


c 












I 0.8 - 




T 












1 




1 


JL, 


1 




T 


1 


■D 








































> 








































■a 

£ 0.6 - 


































1 


















































o 
























































































0) 












































| 0.4 - 












































3 












































z 












































0.2 - 












































0.0 ■ 














































KUR MID PHR LIS LAY MAR GAR FFS MMM NIH 



Figure 5.25. Mean number of individuals per transect from REA data from 
2000-2002 and 2007. Error bars are standard error of the mean. Source: 
NWHI RAMP, unpub. data. 



1.4 

1.2 

| 1.0 

£ 0.8 

V) 

| 0.6 

o 

5 

0.4 

0.2 
0.0 



F 9409 = 3.674 
p < 0.001 



JL 



X 1 



I 



I 



KUR MID PHR LIS LAY MAR GAR FFS MMM NIH 



Figure 5.26. Mean biomass in t ha-1 (ln[x+l]) per transect from REA data 
from 2000-2002 and 2007. Error bars are standard error of the mean. 
Source: NWHI RAMP, unpub. data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 5.14. Rank values for fish assemblage metrics 
gent reefs of the NWHI. Source: NWHI RAMP, unput 


among the 10 emer- 
. data. 


Reef 


m 

3 
Q. 

n 

3 
«>' 

3 


CD Kt 

2 

5' 
(A 


* 

to 

■o 

CD 
O 

CD" 

</> 


II 

9) 
CO 


s-5 1 

9> 
U) 
CO 

* 


Apex 

Predator 

Biomass 


33S 

CD (D 
O J) 

s = 

3 

CD 

3 


33> 

2 £ 

3 CD 

(Q 

CD 


Pearl and Hermes 


10 


9 


4 


9 


10 


10 


6 


8.29 


Midway 


7 


8 


6 


10 


9 


3 


8 


7.29 


French Frigate Shoals 


5 


10 


7 


7 


8 


8 


5 


7.14 


Lisianski 


9 


2 


5 


8 


4 


9 


10 


6.71 


Gardner 


3 


1 


9 


6 


7 


6 


3 


5 


Laysan 


4 


5 


8 


4 


5 


7 


1 


4.86 


Kure 


8 


7 


3 


5 


2 


1 


7 


4.71 


Nihoa 


1 


4 


10 


3 


6 


2 


4 


4.29 


Maro 


6 


6 


2 


2 


3 


5 


2 


3.71 


Mokumanamana 


2 


3 


1 


1 


1 


4 


9 


3 


*Total biomass excludes back reef and 
pares habitat types. 


lagoon habitats to reduce bias and 


com- 



Comparisons of fish assemblage char- 
acteristics among reefs in the NWHI re- 
vealed Pearl and Hermes Atoll to have 
the highest average rank among the 
seven metrics examined (Table 5.14, 
Figure 5.27). Pearl and Hermes yield- 
ed the highest endemism, highest total 
biomass, and highest apex predator 
biomass among all reefs. Midway and 
French Frigate Shoals was second and 
third highest rank with Midway having 
the greatest number of individuals and 
French Frigate Shoals having the high- 
est richness. Lisianski-Neva Shoals had 
the highest recruit index (ratio of YOY to 
older sized individuals). 

Mokumanamana had the lowest rank 
integrated over all fish assemblage met- 
rics and had the lowest number of spe- 
cies per transect and the lowest number 
of individuals per transect. Maro Reef 
and Kure Atoll also had low values for 
most fish assemblage metrics. Since 
all sampling was conducted within the 
boundaries of each Special Preserva- 
tion Area (SPA), these rankings by reef 
should also serve as a ranking by SPA. 



Comparisons with the MHI 
The most conspicuous biological pat- 
terns observed in the NWHI was the 
strikingly higher numerical and biomass 
densities and greater average body siz- 
es of reef fishes in the NWHI compared 
to the MHI, particularly for large jacks, 
reef sharks and other apex predators 

(Figure 5.28). Also notable is the overall reduced numbers and biomass density of lower trophic level fishes 
in the MHI, including lower-level carnivores. Differences in fish biomass density between the MHI and NWHI 
represent both the severe depletion of apex predators from fishing and the heavy exploitation of other species, 
primarily lower trophic-level carnivores on shallow reefs of the MHI (Friedlander and DeMartini, 2002). Fish 
densities at less exploited sites (such as uninhabited Kahoolawe and no-take areas) within the MHI further 
reinforce the conclusions that these differences are caused by fishing. Recent comparisons of fish biomass 
and size structure among accessible sites and inaccessible sites near versus distant from population centers 
in the MHI further indicate that depressed MHI stocks are primarily the result of fishing rather than other an- 
thropogenic stressors such as poorer habitat quality (Williams et al., 2008). Were it not for extraction, reef fish 
productivity in the MHI should be higher (not lower) than in the NWHI as a result of greater terrigenous nutrient 
input and more diverse juvenile nursery habitats at the vegetated, high windward islands. Other anthropogenic 
stressors insufficiently explain the lower densities of reef fishes in the MHI (Friedlander and DeMartini, 2002; 
Friedlander and Brown, 2004). The differences in fish assemblage structure provide evidence of the high level 
of exploitation in the MHI. Further, the sharp contrast between the two areas in terms of fish density and com- 
position provides a valuable perspective for developing ecosystem-level management of reef systems in the 
MHI and the NWHI (Friedlander and DeMartini, 2002). 




Figure 5.27. Mean rank values for fish assemblage metrics among the 10 
emergent reefs of the NWHI. Source: NWHI RAMP, unpub. data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Catalogue of the NWHI Fish Assemblages 

We conclude this chapter with a char- 
acterization of the fish assemblages at 
each of the 10 NWHI reefs, ordered from 
from Nihoa Island to Kure Atoll. The as- 
semblages at each reef are described 
in terms of three basic metrics (species 
richness and numerical and biomass 
densities), with the latter two metrics 
examined for dominant species. 



Nihoa Island 

Despite its small size (Figure 5.29), 
Nihoa Island ranked first overall in fish 
species richness per transect among all 
reefs surveyed in the NWHI. This is in 
contrast to the total species richness, 
which ranked amongst the lowest in the 
NWHI. High species richness is related 
to the proximity to MHI and our obser- 
vation of the highest percentage of spe- 
cies with a tropical-biased distribution. 
Species richness ranged from to 36.6 
to 8.6 ( x =22.8, SD ± 11.02; Table 5.15, 
Figure 5.30). 

Numerical abundance of fishes ranked 
eighth overall and ranged from 5.32 to 
0.14 individuals nr 2 (x =1.61, SD ± 1.57; 
Table 5.15, Figure 5.30). The blackfin 
chromis (Chromis vanderbilti), a plank- 
tivorous damselfish, comprised 35% of 
the total numerical density, followed by 
chubs (16%), and the brown surgeon- 
fish (Acanthurus nigrofuscus, 6%). The 
highest species richness, biomass, and 
numerical abundance were observed 
off the leeward side of the island where 
high complexity basalt benches provid- 
ed good quality habitat for a diversity of 
species of various sizes. 

Biomass ranked fifth overall. Mean bio- 
mass per station was 2.88 t ha 1 (SD ± 
3.37) and ranged from a high of 12.03 to 
a low of 0.39. Chubs accounted for 43% 
of the total biomass at Nihoa Island, fol- 
lowed by whitetip reef sharks (7%), the 
introduced blueline snapper (Lutjanus 
kasmira, 7%) and black durgons (Melichthys niger, 5.5%). 




Figure 5.28. Comparisons of total biomass and biomass among consumer 
groups between the NWHI and MHI. Source: Friedlander and DeMartini, 
2002. 




Figure 5.29. Aerial image of Nihoa Island. Photo: J. Maragos. 

Table 5.15. Fish assemblage characteristics for Nihoa Island. Source: 
NWHI RAMP, unpub. data 



LEVEL 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


11 


22.8 


11.02 


3.32 


15.4 


30.21 


Number of 
Individuals (m 2 ) 


11 


1.61 


1.57 


0.47 


0.55 


2.66 


Biomass (t ha -1 ) 


11 


2.88 


3.37 


1.01 


0.62 


5.14 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Fish Species Richness 


• 1.3-15.0 




• 15 1 • 19.3 




• 19.4-23.3 




23.4-27.0 




£ 27.1-46.7 
1 L*od 


N 
i 


WHw20m 


k 


0.2 04, 






Number of Individuals 


■ 0.0 - 0.B 




• 0.9-1.2 




• 1.3-1.5 




# 1.6-2.1 




% 2,2 ■ 12.0 


N 
j 


| Land 


WHH^Dm 


A 


0.2 04 

^^^^m iKilomoiera j 




o 

o 



Fish Biomass 




* 0,0 - O.S 




o 0.9 ■ 1.3 




O i .i 2.0 




O 21-3.3 




O 3-4-22.9 

JLttd 
W«(*<Zt>i*i 


N 

A 


0.2 0.4 

^^^^^^^=3 Ki kxnerl Era 




J 



Figure 5.30. Fish assemblage characteristics for Nihoa Island. Species richness (top left), number of individuals (top 
right), and biomass (t ha 1 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 



Table 5.16. Fish assemblage characteristics for Mokumanamana Island. 
Source: NWHI RAMP, unpub. data. 



LEVEL 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 
MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


15 


17.82 


4.45 


1.15 


15.36 


20.29 


Number of 
Individuals (m 2 ) 


15 


0.81 


0.35 


0.09 


0.61 


1 


Biomass (t ha 1 ) 


15 


1.25 


1.06 


0.27 


0.67 


1.84 



Mokumanamana 

Mokumanamana yielded the lowest 
species richness per transect (x =17.8, 
SD ± 4.45) among all reefs (Table 5.16, 
Figure 5.31, SD ± 4.45). Higher species 
richness was observed on the north- 
western portion of the island. 

Mokumanamana also had the lowest 
numerical density of fishes observed 

among reefs in the NWHI (0.81 individuals/m 2 , SD±0.35) and ranged from 1.49 to 0.43. The planktivorous 
blackfin chromis accounted for 19% of total numerical density, followed by the saddle wrasse (Thalassoma 
duperrey,18%) and the orangeband surgeonfish (Acanthurus olivaceus, 10%). The low overall values for fish 
assemblage characteristics at Mokumanamana are likely the result of low habitat complexity where the ma- 
jority of stations having extremely low relief. For example, Shark Bay, located on the northern portion of the 
island, exhibited substrate of flat planed surfaces as a result of scouring by surge and sediment suspension. 

Fish biomass also ranked lowest at Mokumanamana. The distribution of biomass was extremely variable (CV 
= 0.85) but was highest off the points on the north and eastern parts of the island. Biomass ranged from 4.11 
to 0.36 t ha 1 with a grand mean of 1.25 (SD ± 1.06). Apex predators accounted for 43% of the total biomass 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Fish 


Sp*C 


&Z RlCrtrWSi 


- 


1.3- 


4.3 




* 


14.3 


19 




• 


ISO 


23.0 




• 


23.0 


2S.6 




• 


26.6 


4&.& 




[_ 


| ; -H---1 

Wai*"tf0m 


A 


a 


0.2 


0.4 





IM'<I'M"W 



164 b aZ , 3&' , VV 




itwrww 



164'« , 36"W 



l64"*rZ<"W 




i 0.1-0.7 

o 0.7-1.2 
O 1.2- IS 

81.9-3.2 
3.2 - 22.9 



tW^'H-W 



Figure 5.31. Fish assemblage characteristics for Mokumanamana. Species richness (top left), number of individuals (top 
right), and biomass (t ha 1 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 



and were dominated by grey reef sharks with 25% of total fish biomass. Other important contributors to fish 
biomass included orangeband surgeonfish (17%), black durgons (17%), giant trevally (6%) and whitetip reef 
sharks (5%). 



French Frigate Shoals Fish 
Species richness at French Frigate 
Shoals averaged 21.8 (SD ± 7.7) and 
was the forth highest among all reefs 
surveyed (Table 5.17, Figure 5.32). Fore 
reef habitats had the highest species 
richness (x= 26.1), followed by back 
reef (x = 20.7), and lagoon habitats (x 
= 19.3). Species richness tended to be 
higher on the windward fore reef (Table 
5.18). 



Table 5.17. Fish assemblage characteristics for French Frigate Shoals 
across all habitat types. Source: NWHI RAMP, unpub. data. 



LEVEL 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


93 


21.78 


7.73 


0.8 


20.19 


23.37 


Number of 
Individuals (m 2 ) 


93 


1.69 


1.83 


0.19 


1.31 


2.07 


Biomass (t ha 1 ) 


93 


2.28 


2.22 


0.23 


1.83 


2.74 



Fish density ranged from 12.0 to 0.26 individuals/m 2 and averaged 1.69 (SD ± 1.8). Numerical abundance 
was highest in lagoon habitat (1.77 individuals/m 2 ) and was dominated by the domino damselfish (Dascyllus 
albisella, 9%), saddle wrasse (8%) and goldring surgeonfish (Ctenochaetus strigosus, 7%). Density was low- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Fish Species Richness 


• 


1 a. 


14.3 




• 


14.3 


ISO 




• 


ig.e 


23.0 




• 


23.0 


28 e 




• 


as 


46.6 






WWv 


.:-.'i m 


A 


1.252 5 


■ K ■■ 


6 tare 








■. 6> 



o 
•- 



Number of Individuals 




o c • 


■0.9 




* 


19 


1.2 




• 


1.2 


1.S 




• 


1.9 


2 ! 




• 


■} l 


L'0 






Land 


A 


1252.5 


5 








im KitaTOterS 












r^^x 










Number of Individuals 

■■ Hmh 1183 

■Kim 0.19 

N 
Ha data t, 

1.252.5 5 


V \# JJ 













1G6 2<T>V 166 '6"W '66 12"W 


1W6W 


























^ — ■ ^^^ 










o ° 

o -° 

o ° 

©■ -, o 


















! 

5 


o 
o 
•0 










\ 










© °» 


o 








\ 








S 

5 






i 


" o 
q 

o 

o 

o 



■ 3 




v. 


\ 




); 






Fish Blomass 

• 0,1-0.7 

o 0.7-1.2 
O 1.2-1.9 
O 19-3.2 
O 3.2-22.8 M 


° O o 


Fish Biomass 
^* Low CUM 






=1— ,; 

Walw *2G m /^ 
01H2.S 5 


° S§ 




o 






N 

•*" uo Al 

1.252.5 5 
i^H=i^i^i^ KjkH neiera 






C ::/ 










1BS"24"W 1W1BYI 16G I2W 


<KTW 















Figure 5.32. Fish assemblage characteristics for French Frigate Shoals: species richness (top row), number of individuals 
(middle row), and biomass (t ha 1 , bottom row). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

. .. f , ,_ 2 . , Table 5.18. Fish assemblage characteristics for French Frigate Shoals for 

esi on me rare reer (X - J..&& m ;, wnere eacn mayo/ , habitat type source: NWHI RAMP, unpub. data. 

the blackfin chromis and saddle wrasse 
each accounted for 10% of the total num- 
ber of individuals. Numerical abundance 
on the back reef habitat (x = 1.55) was 
composed of the saddle wrasse (12%), 
the introduced blueline snapper (9%) 
and blackfin chromis (8%). The greatest 
number of individuals was observed at 
stations near Tern Island at the northern 
portion of the atoll and at the southern 
pass near Disappearing Island. 

Fish biomass density (t ha 1 ) was high- 
est on the fore reef (x = 3.08 t ha 1 ) and 
was dominated by giant trevally (26%) 
and grey reef sharks (21%). French 
Frigate Shoals ranked third in total fore 
reef biomass among all locations. Spe- 
cies composition by weight in the lagoon ( x = 1.87) primarily consisted of giant trevally (13%), followed by grey 
reef sharks (6%), the endemic spectacled parrotfish (6%) and bluespine unicornfish (Naso unicornis, 6%). The 
back reef habitat yielded the lowest biomass (x = 0.97) where giant trevally (25%), bluelined snappers (14%) 
and grey reef sharks (10%) comprised nearly half of the total biomass. Biomass was highest near Tern Island 
at the northern portion of the atoll and at the southern pass near Disappearing Island. 



BACK REEF 


NUMBER 


MEAN 


STD 
DEV 


STD 

ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


2 


20.67 


0.47 


0.33 


16.43 


24.9 


Number of 
Individuals (m 2 ) 


2 


1.55 


0.68 


0.48 


-4.52 


7.61 


Biomass (t ha' 1 ) 


2 


0.97 


0.1 


0.07 


0.06 


1.87 


LAGOON 


Species 


58 


19.34 


6.83 


0.9 


17.54 


21.14 


Number of 
Individuals (m 2 ) 


58 


1.77 


2.16 


0.28 


1.2 


2.34 


Biomass (t ha' 1 ) 


58 


1.87 


1.72 


0.23 


1.42 


2.33 


FORE REEF 


Species 


33 


26.13 


7.63 


1.33 


23.43 


28.84 


Number of 
Individuals (m 2 ) 


33 


1.56 


1.12 


0.19 


1.16 


1.96 


Biomass (t ha -1 ) 


33 


3.08 


2.8 


0.49 


2.09 


4.08 



Table 5.19. Fish assemblage characteristics for Gardner Pinnacles across 
all habitat types. Source: NWHI RAMP, unpub. data. 



NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


10 


22.55 


6.48 


2.05 


17.91 


27.19 


10 


1.39 


0.72 


0.23 


0.88 


1.91 


10 


2.99 


2.11 


0.67 


1.48 


4.5 



Gardner Pinnacles 

A total of 10 stations were sampled for 
fishes at Gardner Pinnacles with the 
sampling effort representing a large por- 
tion of the hard bottom habitat less than 
18.2 m in depth including windward and 
leeward exposures. Mean species rich- 
ness was 22.5 (SD ± 6.5) with a range 
from 36 to 13.3 species per transect. 
Gardner ranked second in mean spe- 
cies richness even though total species richness was low (Table 5.19 and Figure 5.33). 

Despite its small size, the biomass density of fishes at Gardner Pinnacles ranked fourth overall. Fish biomass 
ranged from 7.68 to 0. 29 t ha-1 (* = 2.99, SD ± 2.11; Table 5.19, Figure 5.33). Chubs dominated by weight, 
comprising 17% of the total fish biomass. This species was followed by bluefin trevally (Caranx melampygus, 
11%) and grey reef sharks (10%). Highest biomass was observed off the northwest basalt pinnacle where 
large boulders formed a highly complex habitat with a vertical wall down to the reef pavement at 18.2 m. This 
station was dominated by bluefin (25%) and giant trevally (13%). 

Fish density ranged from 2.88 to 0.62 individuals/m 2 (x = 1.39, SD ± 0.73). Chubs accounted for 16% of total 
numerical abundance, followed by saddle wrasse (9%), and oval chromis (Chromis ovalis, 7%). The drop-off 
at the northwest sea stack harbored a large number of planktivores including oval chromis and milletseed 
butterflyfish (Chaetodon miliaris). These two species comprised 32% of the numerical density of fishes at this 
station. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Fish Species Richness 


* 


1 |- 


14.3 




• 


14-3 


19.0 




• 


19 


23.0 




• 


23 


26.6 




• 


26 6 


46.6 




1 


i.. n 
.■,..■.- •;• - . 


N 

A 


o at 02 

^^^^=^ K i lomol era 



1WW 


» * 




• 




* 




Number of Individuals 


• 


- 001-09 


■ 0.9-1.2 




• 1 .2 - 1 .6 




• fcB-2.1 




2 1 ■ 12,0 




L^d J 1 




WH**<20m f\ 




0.1 02 




^^^^=^ Xi kjrnnl era 





° o O 

o 

8 ° 






o . 


Fish Biomass 
■ 0,1-0.7 
o 0.7-1.2 
O 1.2- 1.9 

O 1.0-3.2 

O 3.2-22.9 

Zi-> ; 

W»W<H>iti /\ 
0,1 02 











Figure 5.33. Fish assemblage characteristics for Gardner Pinnacles. Species richness (top left), number of individuals 
(top right), and biomass (t ha 1 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 



LEVEL 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


42 


18.65 


4.58 


0.71 


17.22 


20.08 


Number of 
Individuals (m 2 ) 


42 


1.26 


0.51 


0.08 


1.1 


1.42 


Biomass (t ha 1 ) 


42 


1.92 


1.7 


0.26 


1.39 


2.46 



Maro Reef 

Maro Reef was second only to Moku- Table 5.20. Fish assemblage characteristics for Maro Reef across all habi- 
manamana Island in having the lowest far types. Source: NWHI RAMP, unpub. data. 
mean species richness observed on 
quantitative surveys (Table 5.20, Figure 
5.34). Mean species richness was 18.6 
(SD ± 4.6) and ranged from 27.67 to 
12.56 per station. Relatively high spe- 
cies numbers were recorded at stations 
along the westernmost, leeward reef 
sections. 

The number of individual fish observed on transects at Maro was also low compared with other reefs in the 
NWHI (x = 1.3, SD ± 0.5). Small resident species such as saddle wrasse (16%), Pacific Gregory (Stegastes 
fasciolatus, 13%) and juvenile parrotfishes (11%) comprised much of the numerical density observed at Maro. 
Several stations on the windward, northeast side of the reef possessed higher numbers of individuals com- 
pared to other stations and were dominated by small juvenile parrotfishes. 

Biomass also was low compared to most other locations and ranked second lowest after Mokumanamana Is- 
land. Biomass density ranged from 6.12 to 0.41 1 m 2 (x = 1.9, SD ± 1.7); and similar to the observed richness 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 





• 

• 

• 

• 


FJsh Specks Richness 

• 1.3-14.3 

• 14.3- 19,0 

• 19.0- 23.D 

• 23,0 -26.6 

% 26 B - 48.6 

mt " A 

Water ^Oni /\ 


• • 

• 











ft 

m 

m 

•• • # • 

• 

• • 

• 


NumWr or Individuals 

• 0.01-05 

• 0.9-1.2 

• 1.2- 1.6 

• 1.0-2.1 

• 2.1 - 12 D 

rn- ! 

WdBr<20m /\ 
1 a A 


• • 

• 











O 

o 



<p 
o 

O ■ o 

■ ° o 
o 


Fish BiWI!H£B 
- 01-07 
0.7 -1.2 
O 12-1.0 
O 19-32 
O 3.2 - 22.8 

rn- I 

o i a *_ 




pes 







Figure 5.34. Fish assemblage characteristics for Maro Reef. Species richness (top left), number of individuals (top right), 
and biomass (t ha 1 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 

patterns, relatively high biomass was observed along the westernmost, leeward reef sections. More than 25% 
of the biomass at Maro Reef consisted of giant trevally, followed by spectacled parrotfish (11%), Galapagos 
sharks (7%), bullethead parrotfish (Chlorurus sordidus, 6%) and whitetip reef sharks (6%). 



Table 5.21. Fish assemblage characteristics for Laysan Island across all 
habitat types. Source: NWHI RAMP, unpub. data. 



Laysan Island Fish 

Laysan Island ranked third in mean spe- 
cies richness ( * = 22.4, SD ± 5.2; Table 
5.21, Figure 5.35), ranging from 32.7 to 
13.3. The highest species richness oc- 
curred on the windward fore reef, off the 
northeast corner of the island. 

Numerical abundance ranged from 2.46 

to 0.42 individuals nr 2 (x= 1.3, SD ± 

0.5) and was dominated by saddle wrasses (18%), followed by convict tangs (Acanthurus triostegus, 11%), 

and Pacific Gregory (7%), respectively. No strong spatial patterns to numerical abundance were observed 

among the sampling stations at Laysan. 



LEVEL 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


20 


22.37 


5.19 


1.16 


19.94 


24.79 


Number of 
Individuals (m 2 ) 


20 


1.34 


0.53 


0.12 


1.1 


1.59 


Biomass (t ha 1 ) 


20 


2.55 


2.09 


0.47 


1.58 


3.53 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 







Fish S|»cie& Richness 


• 


1 3- 


14.3. 




• 


14.3 


19.0 




• 


19.Q 


2&0 




• 


23.0 


2B£ 




• 


26.6 


■16.6 




■= 




N 

A 


• 


05 







. 


• 


■ 






V 












. 


• 








■ 


Number at individual* 




• • 






• 0.01 ■ OS 

* 09-12 










m ia-16 








• l.B-2.1 










2.1 -12.0 
04 1 










Figure 5.35. Fish assemblage characteristics for Laysan Island. Species richness (top left), number of individuals (top 
right), and biomass (t ha 1 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 

Laysan Island ranked sixth in mean biomass (x = 2.5, SD ± 2.1), ranging from 5.15 to 0.85 t ha 1 . The wind- 
ward, northeast fore reef harbored the highest biomass. Giant trevally comprised 37% of total biomass, fol- 
lowed by whitebar surgeonfish (Acanthurus leucopareius, 6%) and the endemic spectacled parrotfish (6%). 



Table 5.22. Fish assemblage characteristics for Lisianski Island-Neva 
Shoals across all habitat types. Source: NWHI RAMP, unpub. data. 



Lisianski Island-Neva Shoals 
Lisianski Island-Neva Shoals ranked 
sixth in mean species richness per sta- 
tion (x = 21.2, SD ± 4.13) and ranged 
from 24.7 to 20.7 (Table 5.22, Figure 
5.36). The greatest number of species 
per station was observed on the leeward 
side (northwest and west) of Lisianski. 

Numerical abundance at Lisianski Island-Neva Shoals ranked third overall. Mean fish density was 1.45 indi- 
viduals/m 2 (SD ± 0.5) and ranged from 1.66 to 0.99 individuals/m 2 . Dominant species include saddle wrasse 
(12%), goldring surgeonfish (11%), Pacific Gregory (11%) and juvenile parrotfish (8%). No spatial patterns 
were observed for fish density across the reef system. 



LEVEL 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


32 


21.23 


4.13 


0.73 


19.75 


22.72 


Number of 
Individuals (m 2 ) 


32 


1.45 


0.53 


0.09 


1.26 


1.64 


Biomass (t ha -1 ) 


32 


2.47 


1.71 


0.3 


1.85 


3.08 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



< 


. • 




• • 


• • 




• 


• 




• 


• 
• 




• 


• 






• 


FlSh SptClfrft Pi; I'll- , ■ 




■ 1.3-14.3 






* 14.3-19.0 






• 1S.Q-Z3.D 






9 23.0-2B.B 




• • 

•*• • • 

• • 


# Sofc-JHR 

Wswrzum /\ 
12 4 







* 








• 










•• 


• 








• - 


• 








o 


• 
• 
• 








• 


• 
• 








• 



















• 


Number of Individuals 

• o.oi - o.a 


' 








* 0.9-1.2 








'J 


• 12-16 

• 1.6-2.1 






□ 




# 2.1 ^ 12 D 




9 * • 


o 




WflW«2Di» f\ 






o 








• • 




lb 









?" 


1M'W 113'54-W 


ITJ*4SVM 


IS 


o 
O 

°° ° 

O o 

O o 






I' 


° 






o 

n ° 

o °o Q o 

O O 


Fish Biomass 
- 0.1- 0.7 
o 0.7-1.2 
O 12-1.8 
O 1.9-3.2 
O 3.2-22.9 

(IS 4 







Figure 5.36. Fish assemblage characteristics for Lisianski Island-Neva Shoals. Species richness (top left), number of 
individuals (top right), and biomass (t ha 1 ; bottom left). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 

Lisianski Island-Neva Shoals ranked seventh in mean biomass compared to all other reefs. Mean fish biomass 
was 2.5 t ha 1 (SD ±1.7) and ranged from 5.15 to 0.8. Biomass was highest along the southeast portion of 
Neva Shoals, in an area of high coral cover and high habitat complexity. Giant trevally accounted for the major- 
ity (51%) of the total biomass. This was followed in importance by three species of parrotfishes: the endemic 
spectacled parrotfish (8%), bullethead parrotfish (4%) and the endemic regal parrotfish (Scarus dubius, 4%). 



Pearl and Hermes Atoll 
Mean species richness at Pearl and 
Hermes Atoll was 20.1 (SD ± 7.2) and 
ranked seventh overall (Table 5.23, Fig- 
ure 5.37). Species richness was signifi- 
cantly higher (F 291 = 24.49, p<0.001) on 
the fore reef (x = 24.0, SD ± 6.9) com- 
pared with the lagoon (x = 16.9, SD ± 
6.0) and back reef (x = 17.0, SD ± 4.6) 
habitats (Table 5.24). 



Table 5.23 Fish assemblage characteristics for Pearl and Hermes Atoll 
across all habitat types. Source: NWHI RAMP, unpub. data. 



LEVEL 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


91 


20.12 


7.21 


0.76 


18.62 


21.62 


Number of 
Individuals (m 2 ) 


91 


1.82 


0.83 


0.09 


1.65 


1.99 


Biomass (t ha -1 ) 


91 


2.78 


3.75 


0.39 


2 


3.56 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



••• - 



•*». 



Fish sp#cie* Riehntss 


- 


1.3- 


14.2 




* 


1.13 


19.0 




• 


ISO 


23.0 




• 


23 il 


26 6 




• 


26 6 


■4B.B 




1 ll~i 




A 




ttwor<0ni 


■:i 1.; 


szs 


V 














17G°6'W 


17G°W 175°54'W 175°48"W 


z 

t. 

z 


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Figure 5.37. Fish assemblage characteristics for Pearl and Hermes Atoll. Species richness (top row), number of individu- 
als (middle row) and biomass (t ha 1 , bottom row). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

The numerical density of fishes at Pearl Table 5.24. Fish assemblage characteristics for Pearl and Hermes Atoll for 
and Hermes ranked second overall ( x each major habitat ty P e - Source: NWHI RAMP ' un P ub ' data - 
= 1.8, SD ± 0.8). Overall, planktivores 
comprised a third (37%) of total den- 
sity, which included oval chromis (11%) 
blackfin chromis (7%) and chocolate 
dip chromis (Chromis hanui, 5%). The 
number of individuals observed on the 
fore reef (x = 2.1, SD ± 0.9) was sig- 
nificantly higher (p<0.05) than the back 
reef (x = 1.9, SD ± 0.6) and lagoon (x 
= 1.6, SD ± 0.8) habitats. 

Pearl and Hermes ranked first in fish 

biomass on the fore reefs (x = 3.9, 

SD± 4.4) among all locations. Biomass 

was significantly higher on the fore reef 

(p < 0.05) than the lagoon (x = 2.0, SD 

± 3.0), which, in tern, was significantly higher (p < 0.05) than at back reef (x = 1.0, SD± 0.9). Apex predators 

dominated the fish biomass, with giant trevally accounting for 48%, followed by whitetip reef sharks (6%), and 

Galapagos sharks (5%). Stations with the highest biomass were located along the leeward, southwest fore 

reef. 



BACK REEF 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


7 


17 


4.59 


1.74 


12.75 


21.25 


Number of 
Individuals (m 2 ) 


7 


1.88 


0.55 


0.21 


1.37 


2.39 


Biomass (t ha 1 ) 


7 


1.05 


0.85 


0.32 


0.26 


1.83 


LAGOON 


Species 


43 


16.91 


5.99 


0.91 


15.07 


18.76 


Number of 
Individuals (m 2 ) 


43 


1.63 


0.78 


0.12 


1.39 


1.87 


Biomass (t ha -1 ) 


43 


2.02 


3.02 


0.46 


1.09 


2.95 


FORE REEF 


Species 


41 


24.02 


6.91 


1.08 


21.84 


26.2 


Number of 
Individuals (m 2 ) 


41 


2.01 


0.88 


0.14 


1.73 


2.29 


Biomass (t ha 1 ) 


41 


3.88 


4.42 


0.69 


2.49 


5.28 



Midway Atoll 

Midway ranked fifth in species richness 
among all reef locations (Table 5.25, 
Figure 5.38). The mean number of spe- 
cies per transect differed significantly 
(p<0.05) among all three habitats. Fore 
reef habitats harbored 27.0 (SD ± 6.0) 
species, followed by back reefs (x = 
19.1, SD ± 4.8), and lagoon habitats (x 
= 15.8, SD ± 6.1). Richness was high- 
est along the southern fore reef (Table 
5.26). 

Midway ranked first in numerical density 
(x =2.7 individuals/m 2 , SD ± 2.0). The 
lagoon harbored the greatest number 
of individuals (x =2.9 individuals/m 2 , 
SD ± 2.7) consisting of damselfishes 
(oval chromis - 11%, Pacific Gregory - 
10%, domino damselfish - 5%, blackfin 
chromis - 5% and chocolate dip chromis 
- 5%). All except the Pacific Gregory 
are planktivores. Saddle wrasse (13%) 
and schools of convict tangs (6%) also 
contributed to the large number of indi- 
viduals observed at Midway. Numerical 
abundance was highest on the north- 
western leeward fore reef and in Welles 
Harbor. 



Table. 5. 25. Fish assemblage characteristics for Midway Atoll across all 
habitat types. Source: NWHI RAMP, unpub. data. 



LEVEL 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


37 


21.59 


7.97 


1.31 


18.94 


24.25 


Number of 
Individuals (m 2 ) 


37 


2.69 


2.04 


0.34 


2.01 


3.37 


Biomass (t ha -1 ) 


37 


2.5 


2.14 


0.35 


1.78 


3.21 



Table 5.26. Fish assemblage characteristics for Midway Atoll for each 
major habitat type. Source: NWHI RAMP, unpub. data. 


BACK REEF 


NUMBER 


MEAN 


STD 
DEV 


STD 
ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


4 


19.08 


4.79 


2.4 


11.45 


26.71 


Number of 
Individuals (m 2 ) 


4 


1.59 


0.46 


0.23 


0.85 


2.32 


Biomass (t ha 1 ) 


4 


1.46 


0.65 


0.33 


0.42 


2.49 


LAGOON 


Species 


15 


15.76 


6.14 


1.59 


12.35 


19.16 


Number of 
Individuals (m 2 ) 


15 


2.9 


2.69 


0.69 


1.41 


4.39 


Biomass (t ha - 1 ) 


15 


1.58 


1.22 


0.31 


0.91 


2.26 


FORE REEF 


Species 


18 


27.02 


6.05 


1.43 


24.01 


30.03 


Number of 
Individuals (m 2 ) 


18 


2.76 


1.58 


0.37 


1.97 


3.54 


Biomass (t ha 1 ) 


18 


3.49 


2.52 


0.59 


2.24 


4.74 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 









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• • 


• 

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• 


• 

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• 


• 




• 








• 




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• 1.6-2.1 




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O 
OP 


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o 






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r 


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o 




o 


o 








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m H> e>> ww 

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Figure 5.38. Fish assemblage characteristics for Midway Atoll. Species richness (top row), number of individuals (middle 
row) and biomass (t ha 1 ; bottom row). Source: NWHI RAMP, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Biomassat Midway averaged 2.5tha 1 (SD±2.1) and ranked second overall among locations. There were large 
differences in biomass among habitat types with biomass on the fore reef (x - 3.5, SD ± 2.5) more than two 
times higher than the back reef (x - 1.5, SD ± 0.7), and the lagoon (x - 1.6, SD ± 1.2). Herbivores accounted 
for the majority of the biomass (57%) with considerable contributions from the spectacled parrotfish (13%), 
whitebar surgeonfish (8%), convict tang (7%), and bluespine unicornfish (6%). Galapagos sharks (8% ) and gi- 
ant trevally (5%) were the major predators by weight. The highest biomass was observed along the northwest 
fore reef where the reef crest becomes submerged and along the southern fore reef off Sand Island. 



LEVEL 


NUMBER 


MEAN 


STD 
DEV 


STD ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


59 


19.6 


6.32 


0.82 


17.96 


21.25 


Number of 
Individuals (m 2 ) 


59 


1.41 


0.81 


0.1 


1.2 


1.62 


Biomass (t ha -1 ) 


59 


1.22 


1.13 


0.15 


0.93 


1.51 



Ku re Atoll 

Species richness at Kure was low (x Table 5.27. Fish assemblage characteristics for Kure Atoll across all habi- 

- 19.6, SD ± 6.3) ranking eighth overall tat types. Source: NWHI RAMP, unpub. data. 
(Table 5.27, Figure 5.39). Significantly 
higher (p<0.05) numbers of species 
were observed on the fore reef (x = 
21.5, SD ± 6.3) compared to the lagoon 
(x = 17.4, SD ± 6.0) and back reef (x 

- 15.3, SD ± 2.0). Richness was high 

around the entire fore reef (Table 5.28). Tabje 52 s. Fish assemblage characteristics for Kure Atoll for each major 

habitat type. Source: NWHI RAMP, unpub. data. 
An average of 1.4 individuals/m 2 were 
observed at Kure (sixth overall). Saddle 
wrasse (20%), oval chromis (12%), Pa- 
cific Gregory (9%) and chubs (6%) were 
most important numerically. No strong 
patterns in the distribution of individu- 
als was observed and no significant dif- 
ference among habitat types (p>0.05) 
were detected. The fore reef averaged 
1.6 individuals/m 2 , followed by lagoon 

(x = 1.3, SD ± 1.1) and back reef (x 
= 1.1, SD ± 0.2). Fish density was high- 
est on the leeward fore reef and central 

western patch reefs. 

Kure had the second lowest biomass of 

any location (x = 1.2, SD ± 1.1) and the 

lowest proportion of apex predators (16%). Spectacled parrotfish (17%), chubs (10%) and giant trevally (5%) 

were most important by weight. There were no strong patterns in the spatial distribution of biomass. Unlike 

other locations, the lagoon (x = 1.2, SD ± 1.5) and fore reef (x = 1.3, SD ± 0.9) biomass estimates were very 

similar. 



BACK REEF 


NUMBER 


MEAN 


STD 
DEV 


STD 
ERR 

MEAN 


LOWER 

95% 


UPPER 

95% 


Species 


5 


15.27 


2.05 


0.92 


12.73 


17.81 


Number of 
Individuals (m 2 ) 


5 


1.11 


0.21 


0.09 


0.85 


1.37 


Biomass (t ha 1 ) 


5 


0.61 


0.19 


0.09 


0.38 


0.85 


LAGOON 


Species 


20 


17.42 


5.96 


1.33 


14.63 


20.21 


Number of 
Individuals (m 2 ) 


20 


1.31 


1.09 


0.24 


0.8 


1.82 


Biomass (t ha - 1 ) 


20 


1.22 


1.53 


0.34 


0.5 


1.94 


FORE REEF 


Species 


34 


21.53 


6.3 


1.08 


19.33 


23.73 


Number of 
Individuals (m 2 ) 


34 


1.52 


0.65 


0.11 


1.29 


1.75 


Biomass (t ha 1 ) 


34 


1.31 


0.91 


0.16 


0.99 


1.63 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 





• •• 

* 
• 

* * 

• • 


Flih Sp*ct» RFchrwu 

' 1 .3- 14.3 
■ 14.3-19.0 

• 19.Q-23.0 

• 23.0-26.6 

9 26.6-J6.6 N 

Wntor«4Qm Jl 
D D.5 1 2 





• • 



• • 



:•• 



or md.»iawis 

• 0.01-0.9 

• 0.9-1.2 

• 1.2-1.6 

» 1.6-2.1 
2.1-11,0 „ 



D O-fi 1 i^ 



A 



f 



O O 



o 

O 



FishB 
■ 0,1-0,7 
o 0.7-1.2 
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1.9-3.2 
3.2 - 22.9 



8 






□ OJ 1 i 



O 
°.* 
8 



T-O^""^^ o 








_ A 




0.5 1 2, 



Figure 5.39. Fish assemblage characteristics for Kure Atoll, 
row) and biomass (t ha 1 ; bottom row). Source: NWHI RAMP, 



Species richness (top row), number of individuals (middle 
unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



EXISTING DATA GAPS 



Two major issues dominate the management and conservation of reef resources in the NWHI and through- 
out the Hawaiian Archipelago. The dispersal, connectivity, and genetic exchange between reef populations 
of NWHI and MHI organisms is undoubtedly the issue of greatest consequence for future management and 
conservation of reef fish and other resources in the NWHI as well as the MHI. Much recent progress has been 
made obtaining the empirical data needed to begin unraveling patterns of planktonic dispersal and more di- 
rected adult movements of fishes in the NWHI (see the Connectivity and Integrated Ecosystem Studies chap- 
ter of this document). The habitat relations of fishes are arguably the second most important issue to consider 
for fishes in the NWHI. Habitat alterations (sea level rise, warming, acidification) resulting from global climate 
change are expected to be the most significant impacts and likely to occur in the NWHI (Selkoe et al., 2008). 
Although the effects of global warming and coral bleaching on coral reef fishes are of concern worldwide 
(Pratchett et al., 2008a), they are relatively more important in the NWHI where resources are now protected 
from other human impacts by establishment of the Monument. 

The prevalence and dynamics of coral and related substrata (e.g., algal secondary cover) represent the most 
obvious habitat issues for shallow-water reef fishes in the NWHI. Although corals are important as a food 
source only for relatively few, specialized fishes in tropical reef ecosystems including Hawaii (Cole et al. 2008), 
corals provide exceedingly important shelter resources (Caley and St. John, 1996). These shelter resources 
are especially important for the relatively small-bodied and predator-vulnerable juvenile life stages of reef 
fishes, particularly early YOY near the time when they settle from the plankton as "recruits" to benthic popula- 
tions (Jones etal., 2004; DeMartini and Anderson, 2007). Coral shelter is nonetheless also important for larger, 
older juveniles and adults (Beukers and Jones, 1997). 

A comprehensive and systematic characterization of the habitat relations of Hawaiian reef fishes is lacking and 
needed. The only published work to date is limited to finger coral habitat on shallow (10 m) fringing reefs of 
the leeward Big Island and further restricted to the recruits of a suite of summer-recruiting species, primarily 
tangs of the family Acanthuridae (DeMartini and Anderson, 2007). Work is in progress to expand this catalogue 
both taxonomically and across additional habitats, with initial emphasis on the diverse labroids (parrotfishes, 
wrasses) that recruit in spring-summer to very shallow (1-3 m deep) and wave-protected coral rubble habitats. 
Several recent case studies exemplify the need for distinguishing habitat relations between juvenile and adult 
conspecifics (Pratchett et al., 2008b; Wellenreuther and Clements, 2008). For this reason, the habitat relations 
of adult as well as juvenile Hawaiian reef fishes are being described. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Marine Protected Species 

Charles Littnan 1 , Marie (Chapla) Hill 2 , Stacy (Kubis) Hargrove 1 , Kaylene E. Keller 3 and Angela D. Anders 4 




INTRODUCTION 

The Monument protects habitat for 
many marine mammal species .includ- 
ing 24 species of cetaceans that have 
been sighted in the Hawaiian Islands 
Exclusive Economic Zone (EEZ), and 
the critically endangered Hawaiian 
monk seal. (Figure 6.1). Twenty three 
species of cetaceans were observed 
and identified to the species level dur- 
ing a 2002 survey (Barlow et al., 2004). 
Four of these species are on the U.S. 
Endangered Species list, including the 
humpback whale (Megaptera novaean- 
gliae), sperm whale (Physeter macro- 
cephalus), fin whale (Balaenoptera physalus) and sei whale (Balaenoptera borealis; http://www.nmfs.noaa. 
gov/pr/species/esa/mammals.htm). In addition, the false killer whale (Pseudorca crassidens) is listed as a 
strategic stock under the 1994 amendments to the Marine Mammal Protection Act (MMPA). Each of these spe- 
cies has been observed within the Monument boundaries (Barlow et al., 2004; Johnston et al., 2007; NMFS, 
unpublished data). Several of the cetacean species observed within the Hawaiian Islands EEZ are found there 
year-round (e.g., spinner dolphins, false killer whales, rough-toothed dolphins). Others occur there only sea- 
sonally and in some cases are known to migrate long distances to use the area for breeding (e.g., humpback 
whales). 



Figure 6.1. The Monument protects habitat for many marine mammal spe- 
cies, including several species of dolphins and whales. A spinner dolphin 
(left) and humpback whales (right) are pictured here. Photos: PIFSC and 
D. Shapiro. 



In addition to providing important habitat 
to cetaceans, the Monument is also the 
primary habitat for the federally endan- 
gered Hawaiian monk seal (Monachus 
schauinslandi; Figure 6.2J. The six main 
Hawaiian monk seal subpopulations are 
completely contained within the Monu- 
ment boundaries. The Hawaiian monk 
seal population is estimated to have 
declined by 60% since the 1950s (An- 
tonelis et al., 2006), and is currently es- 
timated at just under 1,200 individuals. 
Because of its small population size and 
swift rate of decline, the Hawaiian monk 
seal is the focus of intense conservation 
efforts. 








Figure 6.2. The Monument is home to the six primary Hawaiian monk seal 
subpopulations (left). The area also protects nesting habitat of the green 
turtle (right). The Hawaiian monk seal is listed as endangered and the 
green turtle is listed as threatened under the U.S. Endangered Species Act. 
Photos: J. Watt. 



Finally, the Monument provides the primary nesting habitat for the green turtle (Chelonia mydas; Figure 6.2) 
in the Hawaiian Archipelago, listed as Threatened under the U.S. Endangered Species Act (ESA). Green, 
loggerhead (Caretta caretta), hawksbill (Eretmochelys imbricata), leatherback (Dermochelys coriacea) and 



1. NOAA/NMFS/Pacific Islands Fisheries Science Center 

2. Joint Institute for Marine and Atmospheric Research 

3. NOAA/NOS/ONMS/Papahanaumokuakea Marine National Monument 

4. Clancy Environmental Consultants, Inc. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

olive ridley (Lepidochelys olivacea) turtles use the Monument for foraging habitat and as migration pathways. 
Although the Hawaiian green turtle population has been increasing over the past three decades, individuals 
still nest primarily at French Frigate Shoals in the Northwestern Hawaiian Islands (NWHI). 

CETACEANS 

Available Data 

Most of what is known about cetaceans found within the Hawaiian EEZ comes from data collected within the 
waters surrounding the Main Hawaiian Islands (MHI). Although somewhat limited, information on the occur- 
rence of cetacean species within the Monument comes from historical whaling records, documented oppor- 
tunistic sightings and stranding records, data collected during ship-based surveys, and species-specific (e.g., 
spinner dolphin) photo-identification and genetic research. 

Charles Townsend (1935) used whaling logbooks to record locations of certain whale species onto charts of 
both the Atlantic and Pacific Oceans. The Wildlife Conservation Society digitized the charts and made them 
available to the general public (http://www.wcs.org/sw-high_tech_tools/landscapeecology/townsend_charts). 
Data on sperm whale sightings within the Hawaiian EEZ, including the Monument, are extracted from these 
records (see Species Descriptions section, sperm whale). 

Edward W. Shallenberger (1981) collected information from published and unpublished literature, field notes, 
ships' logs, and interviews with knowledgeable people in order to document the occurrence and status of ce- 
tacean species found in Hawaiian waters, including the NWHI. 

The Atoll Research Bulletin produced a series of publications on the natural history of the NWHI in which re- 
cords of cetacean species were documented from opportunistic sightings and stranding events (Rice, 1960; 
Amerson, 1971; Woodward, 1972; Amerson et al., 1974; Nitta and Henderson, 1993). These records are more 
anecdotal in nature and lack precise location data (i.e., latitude/longitude points), however, they are useful for 
generalizing about the occurrence of certain species within the Monument. 



Precise data regarding the occurrence of 
cetaceans in the Monument come from 
recent ship-based surveys within the 
200 nm EEZ surrounding the Hawaiian 
Islands (Barlow et al., 2004; Johnston 
et al., 2007; NMFS, unpublished data). 
In 2002, Barlow et al. (2004) conducted 
standard ship-based visual line-transect 
surveys, from two ships, for all cetacean 
species in the Hawaiian Islands EEZ. 
Search effort included a total of 24,738 
km (13,357 nm) over 157 survey days. 
Acoustic monitoring, photo-documenta- 
tion, biopsy and behavioral studies were 
conducted concurrently. Line-transect 
surveys resulted in observations of 23 
cetacean species within the Hawaiian 
Islands EEZ, with 15 of those species 
seen within the boundaries of the Monu- 
ment (see Species Descriptions section 
and Figure 6.3; Barlow et al., 2004). 
Abundance estimates and densities per 
1,000 km 2 were calculated for 19 of the 23 cetacean species observed (Table 6.1; see Cetaceans Abundance 
Estimates section on page 210; Barlow, 2006). 




Legend 

* Celacean Observations 

Cetacean Survey Transects 

~ | Main Hawaiian Islands 

I I Papahanaumokuakea Marine National Monumeni Boundary 
800 Kilometers 



Figure 6.3. Cetacean Observations and Survey Transects . Sources: Bar- 
low et al., 2004; Johnston et al., 2007; NMFS, unpublished; Townsend, 
1935; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Table 6.1. Estimated abundance of 19 cetacean species in the MHI and outer Hawaiian Islands EEZ. Overall abundances, 
overall densities, and coefficients of variation (CV) are pooled from the MHI and outer EEZ estimates. Pooled abundance 
and density estimates are given for delphinids and beaked whales. Asterisk (*) indicates a more recent estimate of false 
killer whale abundance in offshore waters of the Hawaiian EEZ (484 individuals, CV=0.93) comes from Barlow and Rankin 
(2007). Source: Barlow, 2006. 



SPECIES 


MAIN ISLAND 

ABUNDANCE 

(n) 


OUTER EEZ 

ABUNDANCE 
(n) 


OVERALL 

ABUNDANCE 

(n) 


OVERALL DENSITY 

PER 1,000 km 2 

(D) 


CV 


Offshore spotted dolphin 


4,283 


4,695 


8,978 


3.66 


0.48 


Striped dolphin 


660 


12,483 


13,143 


5.36 


0.46 


Spinner dolphin 


1,488 


1,863 


3,351 


1.37 


0.74 


Rough-toothed dolphin 


1,713 


6,997 


8,709 


3.55 


0.45 


Bottlenose dolphin 


465 


2,750 


3,215 


1.31 


0.59 


Risso's dolphin 


513 


1,859 


2,372 


0.97 


0.65 


Fraser's dolphin 





10,226 


10,226 


4.17 


1.16 


Melon-headed whale 





2,950 


2,950 


1.20 


1.17 


Pygmy killer whale 


956 





956 


0.39 


0.83 


False killer whale * 





236 


236 


0.10 


1.13 


Short-finned pilot whale 


3,190 


5,680 


8,870 


3.62 


0.38 


Killer whale 





349 


349 


0.14 


0.98 


Sperm whale 


126 


6,793 


6,919 


2.82 


0.81 


Pygmy sperm whale 





7,138 


7,138 


2.91 


1.12 


Dwarf sperm whale 





17,519 


17,519 


7.14 


0.74 


Blainville's beaked whale 





2,872 


2,872 


1.17 


1.25 


Cuvier's beaked whale 





15,242 


15,242 


6.21 


1.43 


Longman's beaked whale 





1,007 


1,007 


0.41 


1.26 


Bryde's whale 





469 


469 


0.19 


0.45 


Delphinids pooled 


13,267 


50,087 


63,354 


25.83 




Beaked whales pooled 


371 


19,121 


19,492 


7.95 





The work of Johnston et al. (2007) focused on visual and acoustic observations of humpback whales from 
Oahu to Midway Atoll in the NWHI during March and April 2007. Search effort covered 1,690 km over 12 sur- 
vey days. Surveys were focused on areas near atolls and islands, and tracklines generally followed the 183 
m (1,000 fathom) isobath, however, some tracklines were in deeper waters (Johnston et al., 2007; NMFS, 
unpublished data). Observations were conducted following standard distance sampling/line transect methods 
for cetaceans, similar to those employed in Barlow et al. (2004). During the surveys, sightings of all cetaceans 
were recorded, including location and group size estimates (NMFS, unpublished data). Additionally, acoustic 
monitoring, photo-documentation and biopsy sampling were conducted. These surveys resulted in the detec- 
tion of 44 groups of eight cetacean species. Seven of these species were observed within Monument boundar- 
ies (Johnston et al., 2007; NMFS, unpublished data). In the Species Description section (below) these surveys 
will be referred to as the NWHI survey. 

Other species-specific cetacean work within the Monument includes research on the social patterns and popu- 
lation structure of spinner dolphins (Stenella longirostris) at Midway Atoll (Karczmarski et al., 1998; 1999; Rick- 
ards et al., 2001; Karczmarski et al., 2005) and the genetic diversity of spinner dolphins within the Hawaiian 
Archipelago (Andrews et al., 2006). The photo-identification and behavioral research at Midway Atoll consisted 
of land and small boat-based surveys conducted in 1998 through 2001, totaling 1,104 effort hours. Results of 
this work indicate that a resident spinner dolphin population is present at Midway, but that movement of indi- 
viduals between Midway, Kure Atoll, and Pearl and Hermes Atoll occurs (Karczmarski etal., 1998; 1999; Rick- 
ards et al., 2001; Karczmarski et al., 2005). The analysis of biopsy samples taken from various locations within 
the Hawaiian Archipelago (e.g., Hawaii, Maui/Lanai, Oahu, Niihau, French Frigate Shoals, Pearl and Hermes 
Atoll, Midway Atoll and Kure Atoll) indicates a genetic distinction among separate spinner dolphin populations 
(Andrews etal., 2006). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Species Descriptions 

Description, range and habitat information is taken from the following sources: Tomich (1986), Ridgeway and 
Harrison (1994), Ridgeway and Harrison (1999), Perrin et al. (2002), Reeves et al. (2002), and OBIS SEAMAP 
(an online data source that compiles information from various published sources, http://seamap.env.duke. 
edu/). 

Offshore Pantropical Spotted Dolphin (Stenella attenuata) 

Three subspecies of the pantropical spotted dolphin are recognized worldwide and differ in body size, color- 
ation and skull characteristics. Their lengths range from 1.6 m to 2.6 m and their average weight is 114 kg. 
In general, the pantropical spotted dolphin has a moderately slender body; a relatively small dorsal fin that 
is strongly falcate (back-curved or shaped like a sickle); and a long, slender beak. The basic color pattern 
includes a dark gray dorsal cape that dips low onto the sides below and forward of the dorsal fin and a lighter 
gray ventral (belly) color that widens along the sides of the peduncle, or tailstock. Adults have a black "mask" 
and a dark jaw-to-flipper stripe. The pattern of spotting and striping on adults can be extremely complex and 
variable; in Hawaii animals have very little dorsal spotting. Calves are born without spots, and juveniles de- 
velop them first on their ventral side. The tip of the beak is white and more conspicuous in some populations, 
including those found in the waters surrounding the Hawaiian Archipelago. 

Pantropical spotted dolphins are found in all tropical to warm temperate oceanic waters between 40°N and 
40°S. In the Pacific Ocean, the stocks are separated into the offshore eastern tropical Pacific, the coastal wa- 
ters between Baja California and the northwestern coast of South America, and the near-shore waters around 
the Hawaiian Islands. 



Pantropical spotted dolphins are found 
throughout the Hawaiian Archipelago 
and are common on the leeward sides 
of the islands as well as at shallow off- 
shore banks (Shallenberger, 1981; To- 
mich, 1986). A series of 12 aerial sur- 
veys (1993-1998) within 46 km of the 
coastline surrounding the MHI resulted 
in observations of 23 groups of spotted 
dolphins, with an average size of 42.8 in- 
dividuals (Mobley et al., 2000). Baird et 
al. (2003) recorded 25 sightings around 
the MHI in May and June of 2003 with 
a mean group size of 77.1 individuals. 
Baird et al. (2006) determined that the 
peak in sighting rates off the MHI oc- 
curred in water depths of 1,000-2,000 
m. Within the Hawaiian EEZ, during the 
2002 (August -November) ship-based 
survey, Barlow et al. (2004) recorded 12 
sightings of spotted dolphins in groups 
as large as 80 individuals (Figure 6.4). One sighting of 74 individuals occurred within the Monument boundar- 
ies (Barlow et al., 2004). 




Figure 6.4. Pantropical spotted dolphin observations from 2002 survey of 
the Hawaiian Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 



Striped Dolphin (Stenella coeruleoalba) 

The striped dolphin reaches a maximum length of 2.7 m and a maximum weight of 160 kg. It has a small to 
medium-sized robust body; a prominent falcate dorsal fin; and a long, well-defined beak. The color pattern is 
a combination of bluish-gray and white. The cape, beak, fins and tail are dark blue-gray The ventral side, or 
belly, is white. On each side of the body a narrow black stripe runs from the beak to the eye and then diverges; 
one branch runs from to the pectoral flipper and the other continues along the side to the anal region. A bluish 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

or light gray shoulder blaze is present above the side stripe and below and anterior to the dorsal fin. The mark- 
ings and boldness of the stripes vary with individual and geographical location. 

Striped dolphins are primarily found in 
tropical and warm temperate waters 
that are oceanic and deep from 50°N 
to 40°S. They occur in the U.S. off the 
west coast, in the northwestern Atlantic, 
in the Gulf of Mexico and in the waters 
off of the Hawaiian Islands. This spe- 
cies associates with upwelling areas 
and convergence zones. 




Striped dolphins are considered rare in 
Hawaiian waters (Shallenberger ,1981; 
Tomich, 1986). Baird et al. (2005a) sug- 
gested that they are likely to "use the 
[Hawaiian] islands seasonally (in warm- 
water periods)." In June 2003, Baird et al. 
(2003) recorded one sighting of striped 
dolphins off of Niihau, in approximately 
2,800 m depth, with a mean group size 
of 45. Barlow et al. (2004) recorded 12 
sightings of striped dolphins within the 
Hawaiian EEZ, between August and 

November, with a mean group size of 12.8 individuals (Figure 6.5). Three of these sightings occurred within 
the Monument boundaries (Barlow et al., 2004). 



Figure 6.5. Striped dolphin observations from 2002 survey of the Hawaiian 
Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 



Spinner Dolphin (Stenella longirostris) 

Four subspecies of spinner dolphins are recognized worldwide and include Stenella longirostris longirostris 
(Hawaiian or Gray's), S.I. orientalis (Eastern), S.I. centroamericana (Central American) and S.I. roseiventris 
(Dwarf). They vary regionally in both form and color pattern. Generally, the spinner dolphin body is slender 
and small, reaching maximum lengths of 2.4 m and 78 kg. The melon is relatively flat and the beak is long and 
slender. The dorsal fin shape varies from moderately falcate to triangular. The color pattern is defined by a dark 
dorsal cape that does not dip low along the sides (like the spotted dolphin, a similar-looking species), lighter 
gray sides, and a light gray or white ventral side. 

Spinner dolphins are found in all tropical and subtropical oceans between 30°N and 30°S. Stenella longirostris 
longirostris is the most widely spread subspecies and is typically found around oceanic islands in the Atlantic, 
Indian, western and central Pacific Oceans. Spinner dolphins associated with the islands in the Hawaiian Ar- 
chipelago often use shallow inshore waters to rest and socialize during the day and move offshore at night to 
feed (Norris and Dohl, 1980). 

Hawaiian spinner dolphins are resident throughout the Hawaiian Islands, including the NWHI (Norris and Dohl, 
1980; Shallenberger, 1981; Tomich, 1986; Karczmarski, 2005). Historical reports demonstrate the presence 
of spinner dolphins in the NWHI at French Frigate Shoals, Pearl and Hermes Atoll and Kure Atoll (Amerson, 
1971; Woodward, 1972; Amerson et al., 1974). According to Amerson et al. (1974), prior to 1968 there had 
been no reported sightings of spinner dolphin at Pearl and Hermes Atoll. Photo-identification research (2006- 
2008) indicates the presence of a resident population at Pearl and Hermes Atoll (NMFS, unpublished data). 
Shallenberger (1981) recorded sightings at Laysan, Lisianski and Maro Reef. However, photo-identification 
and behavioral research at Midway Atoll (1998-2001) demonstrated that a resident population of spinner dol- 
phins was present, but the population size changed from 1998 (n= 260 individuals) to 2001 (n=140 individuals) 
due to the immigration of individuals to Kure Atoll (Rickards et al., 2001; Karczmarski et al., 2005). During the 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



2002 (August - November) Hawaiian 
EEZ survey, eight sightings of spinner 
dolphins were recorded; three of these 
sightings were within the Monument 
boundaries (Barlow et al., 2004; Fig- 
ure 6.6). During the 2007 (March-April) 
NWHI survey, two groups of spinner 
dolphins were observed at Gardener 
Pinnacles (NMFS, unpublished data). 
Genetic data from spinner dolphins in 
the MHI suggest that limited genetic ex- 
change is occurring from one island to 
the next (Andrews et al., 2006; Baird et 
al., 2005a; Galver, 2002). Andrews et al. 
(2006) also found that the spinner dol- 
phin populations in the NWHI (Midway, 
Pearl and Hermes and Kure) are geneti- 
cally homogenous and distinct. 




Legend 

A Stenetla longirostris Observations 

Cetacean Survey Transects (Barlow elal.. 2004) 
| Main Hawaiian Islands f . 

I I Papah3naumoku3J<e3 Marina National Monumant Boundary 

200 400 600 BOO 

iKilometers 



Figure 6.6. Spinner dolphin observations in Hawaiian Island EEZ. Sources: 
Barlow et al., 2004; Johnston et al., 2007; NFMS, unpublished; map: K. 
Keller. 



Rough-toothed Dolphin 
(Steno bredanensis) 

The rough-toothed dolphin has a moderately robust body shape; it reaches a maximum length of 2.7 m and a 
maximum weight of 160 kg. The dorsal fin is tall, moderately falcate and located at mid-back. The pectoral flip- 
pers are relatively large. The head is small and lacks a crease between the melon and the long beak. The color 
pattern is muted but defined by a dark dorsal cape, lighter gray sides and a white belly. Most rough-toothed 
dolphins have a mottled appearance from dark spotting on the sides, throat and belly. White scarring (from 
cookie cutter shark bites or other rough-toothed dolphins) is typical. The upper surface of the beak is dark and 
the lips and ventral surface are white. 



Rough-toothed dolphins occur in deep warm temperate and tropical waters worldwide, between 45°N and 
35°S. 



Rough-toothed dolphins are observed 
in deep water around all of the MHI 
(Shallenberger, 1981; Tomich, 1986; 
Figure 6.7). Research conducted be- 
tween 2000 and 2006 in the MHI (Baird 
et al., in press b) demonstrated that 
rough-toothed dolphins use the area 
year-round and are most commonly 
found in waters greater than 1,500 m 
deep. Within the waters of the NWHI, 
Shallenberger (1981) noted sightings of 
rough-toothed dolphins as far north as 
Mokumanamana. Nitta and Henderson 
(1993) reported a sighting at French 
Frigate Shoals. During the 2002 (August 
- November) Hawaiian EEZ survey, 19 
groups of rough-toothed dolphins were 
observed ranging in size from two to 
15 individuals; three of these sightings 
were within the Monument boundaries 
(Barlow et al., 2004). During the 2007 




Figure 6.7. Rough-toothed dolphin observations in Hawaiian Island EEZ 
Sources: Barlow et al., 2004; Johnston et al., 2007; NFMS, unpublished; 
map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

(March-April) NWHI survey, one group of rough-toothed dolphins was observed near Lisianski Island (NMFS, 
unpublished data). 



Bottlenose Dolphin (Tursiops truncatus) 

The bottlenose dolphin is characterized by coastal and offshore ecotypes that are morphologically distinct, in 
which the offshore animals tend to be larger in size. In general, the bottlenose dolphin has a wide head and 
robust body that reaches a length of 2.5 m to 3.8 m and a maximum weight of 500 kg. Males are typically larger 
than females. The beak is short, and there is a distinct crease where it meets the melon. The pectoral flippers 
are long, and the dorsal fin is moderately tall and falcate. The color pattern ranges from a dark gray dorsal cape 
to lighter gray sides but lacks a distinct demarcation. The belly tends to be off-white or pinkish. 

The bottlenose dolphin occurs worldwide in tropical and temperate waters within the range of 45°N and 45°S. 
Coastal populations are found along continents and around most oceanic islands, where they move into or 
reside in bays, estuaries and lower bodies of rivers. Pelagic, or offshore populations tend to reside far offshore 
as in the Gulf Stream of the North Atlantic and the eastern tropical Pacific. 



Bottlenose dolphins are distributed 
throughout the Hawaiian Archipelago 
and occur regularly in the waters sur- 
rounding the MHI (Shallenberger, 1981; 
Tomich, 1986). Research within the MHI 
suggests that individuals are resident to 
particular islands, not mixing between 
islands (Baird et al., 2003; Baird et al., 
2006; Baird et al., in press a). Sightings 
of dolphins in the MHI occurred in aver- 
age depths of 222 m (Baird et al., 2003). 
Baird et al. (2006) demonstrated that off 
Kauai and Niihau there are two existing 
populations of bottlenose dolphins that 
are distinguished by depth preference, 
however, they are not reproductively 
isolated as individuals move between 
the two populations. Historical reports 
demonstrate the presence of bottlenose 
dolphins in the NWHI at Laysan Island, 
French Frigate Shoals, Kure Atoll, and 
Pearl and Hermes Atoll (Rice, 1960; 

Amerson, 1971; Woodward, 1972;Amerson etal., 1974). During the 2002 (August- November) Hawaiian EEZ 
survey, 14 groups of bottlenose dolphins were observed and ranged in size from 4 to 28 individuals; one of 
these sightings was within Monument boundaries, southeast of Midway Atoll (Barlow et al., 2004; Figure 6.8). 
During the 2007 (March-April) NWHI survey, three groups of bottlenose dolphins were observed near Nihoa, 
Mokumanamana and Laysan Islands (NMFS, unpublished data). 




Legend 

a Tursiops Iruncalus Observations 

Cetacean Survey Transects (Barlow et al , 20Q4) 
B Main Hawaiian Islands f\ 

I Papahanaumofcuakea Marine National Monument Boundary 



Figure 6.8. Bottlenose dolphin observations in the Hawaiian Island EEZ 
Sources: Barlow et al., 2004; Johnston et al., 2007; NMFS, unpublished; 
map: K. Keller. 



Risso's Dolphin (Grampus griseus) 

The Risso's dolphin has a bulbous, beakless head with a distinguishable longitudinal crease along the center 
of the melon. It has a robust body that tapers to a narrow tailstock and reaches a length of 2.6 m to 4 m and a 
weight of 300 to 500 kg. The dorsal fin is tall, erect and moderately falcate; and the pectoral fins are long and 
sickle-shaped. The color pattern can be variable from black, dark gray, brown or white and typically lightens as 
an individual ages. Most adults are heavily scarred with teeth rakes of other dolphins, cookie cutter shark bites 
and circular marks from their prey (e.g., squid). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Risso's dolphins are extensively distrib- 
uted throughout tropical and warm tem- 
perate waters. They are generally found 
in offshore waters deeper than 1,000 m 
and with surface temperatures of 10 to 
28°C. Although migration patterns are 
unknown, seasonal shifts in population 
density are recognized and presumed to 
be related to changes in water tempera- 
ture and prey (e.g., squid) abundance. 

Sightings of Risso's dolphins are rare in 
Hawaiian waters (Shallenberger, 1981; 
Tomich, 1986). During a series of aerial 
surveys (in 1993, 1995 and 1998) in 
the MHI only two individuals were seen 
(Mobley et al., 2000), however, these 
surveys only covered waters within 46 
km (25 nm) of the coast. Stranding re- 
cords from five events also demonstrate 
the presence of Risso's dolphins within 
the MHI (Nitta, 1991; Maldini, 2005). During the 2002 (August-November) Hawaiian EEZ survey, seven groups 
of Risso's dolphins were observed (Figure 6.9). In five of these sightings, other cetacean species were pres- 
ent and included short-finned pilot whales, bottlenose dolphins, sei or Bryde's whales, and unidentified small 
dolphins (Barlow et al., 2004). In October 2002, a group of eight individuals was seen south of Lisianski Island 
(Barlow et al., 2004). 




Legend 

A Grampus grisaus Observations 

Cetacean Survey Transects (Barlow et al-, 20CM) 
^ Main Hawaiian Islands 

| Papananaurnokuakea Marine National Monument Boundary 



Figure 6.9. Risso's dolphin observations from 2002 survey of the Hawaiian 
Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 



Fraser's Dolphin (Lagenodelphis hosei) 

The Fraser's dolphin has a stocky body and grows to a maximum length of 2.7 m and a maximum weight of 

210 kg. It has small appendages and a small, well-defined beak. The dorsal fin is triangular to slightly falcate 

and is more erect in males than females. The color pattern is characterized by a dark, grayish-blue dorsal cape; 

lighter gray sides; a whitish belly; a distinctive dark stripe on each side that runs from the eye to the anus; and 

a dark flipper stripe that merges with the 

side stripe along the lower jaw. 

The Fraser's dolphin is a pantropical 
species that is generally found between 
30°N and 30°S. It is primarily an ocean- 
ic species and found in waters deeper 
than 1,000 m, however, it has also been 
found in areas where deep water ap- 
proaches the coast. 

The first documentation of Fraser's 

dolphins in Hawaiian waters occurred 

during the 2002 Hawaiian EEZ survey 

(Barlow et al., 2004; Figure 6.10). Two Tie*** 

groups, comprised of 47 and 171 indi- sstwm,$mvf«mam^»-i*' 

viduals, were observed within the Ha- . , "»"■"-'""*■» 

. - | | Papahanaumokuakea Marine National Monument Boundary 

waiian EEZ (outside of Monument wa- ^ « »» n» 

^^^ ^^^m 1 1 1 1 1 1 I i 

ters) in November (Barlow et al., 2004). 




Figure 6.10. Fraser's dolphin observations from 2002 survey of the Hawai- 
ian Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Melon-headed Whale (Peponocephala electra) 

The melon-headed whale has a moderately robust body that tapers at both ends. This whale can reach a 
length of 2.7 m and a weight of 275 kg. The head is narrow, triangular shaped, and has no beak. The dorsal fin 
is tall (as much as 30 cm), falcate and positioned at mid back. The general color pattern has an overall gray or 
black appearance with variable lighter gray ventral markings and white lips. The face has a dark mask over the 
eyes, which helps to distinguish this species from others that are similar looking (e.g., pygmy killer and false 
killer whales). 

The melon-headed whale is a pantropical species that is found in deep waters between 40°N and 35°S. 

Melon-headed whales have been ob- 
served somewhat regularly in the wa- 
ters surrounding the MHI (Shallenberg- 
er, 1981; Mobley et al., 2000; Baird et 
al., 2003; Baird et al., 2005a). The first 
record is from Hilo Bay in 1841 when a 
group of 60 animals were driven ashore 
by natives (Tomich, 1986). Shallenberg- 
er (1981) noted that a group of 75-150 
melon-headed whales was seen regu- 
larly off the Kohala coast of Hawaii. Dur- 
ing 12 aerial surveys (within 46 km from 
the coast of the MHI) in 1993-1998, 
Mobley et al. (2000) observed three 
groups with an average of 13.5 individu- 
als per group. Between 2000 and 2005, 
Baird et al. (2005a) observed 18 groups 
with an average of 305 individuals per 
group (in the MHI). Three sightings with- 
in 2003 were in an average water depth 
of 1,100 m (Baird et al., 2003). During 




Figure 6.11. Melon-headed whale observations from 2002 survey of the 
Hawaiian Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 



the 2002 (August-November) Hawaiian EEZ survey, Barlow et al. (2004) observed one group of 171 melon- 
headed whales within the northwestern portion of the EEZ (outside of Monument waters; Figure 6.11). 



Pygmy Killer Whale 
(Feresa attenuata) 
The pygmy killer whale has a moderate- 
ly robust body that tapers more toward 
the back half. It grows to a length of 2.6 
m and a weight of 170 kg. The head is 
rounded with no beak. The dorsal fin is 
tall, falcate and positioned slightly be- 
hind the mid back. The color pattern 
is mostly dark gray to black with white 
markings on the lips and belly. Areas of 
lighter gray extend along the sides from 
the eye to the anus. 

The pygmy killer whale is a pantropical 
species that is found between 40°N and 
35°S. It is one of the most poorly known 
species of odontocetes. 




Legend 
A Feresa attenuata Observations 

Cetacean Survey Transects (Ba rlow et a I . . 2004) 

Main Hawaiian Islands * 

I I Papahanaumokuekea Manne National Monument Boundary 

aoo 
uKilomelers 



Figure 6.12. Pygmy killer whale observations from 2002 survey of the Ha- 
waiian Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Pygmy killer whales have been observed off the leeward coasts of Oahu and Hawaii on numerous occasions 
in groups as large as 100 individuals, but typically in groups smaller than 50 individuals (Shallenberger, 1981; 
Tomich, 1986). Between 2000 and 2005, six groups of pygmy killer whales with an average group size of 11.5 
individuals were observed in the MHI (Baird et al., 2005a). Baird et al. (2003) observed one group of 13 indi- 
viduals off of Kauai/Niihau in 613 m depth. A long-term study of opportunistic sightings of pygmy killer whales 
suggests that a small population of year-round residents is located off the island of Hawaii (McSweeney et al., 
in press) Barlow et al. (2004) observed three groups of pygmy killer whales within the Hawaiian EEZ (Figure 
6.12). One group of five individuals was seen within the Monument boundaries near Midway Atoll (Barlow et 
al., 2004). 

False Killer Whale (Pseudorca crassidens) 

The false killer whale has a relatively slender body that grows to a length 6 m and a weight of 2,000 kg. The 
head is small and conical with the melon overhanging the tip of the lower jaw. The dorsal fin is moderately tall, 
falcate and positioned at mid back. The flippers are broad at the base, narrow at the tip and have a bulge in the 
middle of the leading edge that is a distinguishing feature. The color pattern appears dark gray to black over 
the entire body with lighter gray patches on the throat and chest. 

False killer whales are distributed throughout all tropical and warm-temperate waters, generally occurring be- 
tween 50°N and 50°S. They are typically found in waters deeper than 1,000 m. 



Two populations of false killer whales 
are recognized within the Hawaiian 
EEZ (Chivers et al., 2007). Genetic and 
photographic evidence demonstrate 
the presence of an offshore popula- 
tion closely related to animals found in 
the eastern North Pacific (and Palmyra 
Atoll), and an inshore population that is 
demographically distinct (Chivers et al., 
2007; Baird et al., 2008). Both popula- 
tions are listed as one strategic stock 
under the amendments to the MMPA 
as a result of interactions with Hawaii 
longline fisheries (http://www.nmfs. 
noaa.gov/pr/species/esa/mammals. 
htm). According to Baird et al. (2008), 
false killer whales are observed infre- 
quently in the nearshore waters of the 
MHI. Between 1986 and 2006, only 50 
groups of false killer whales were en- 
countered (Baird et al., 2008). During 
aerial surveys within 46 km of the coast- 
line (1993-1998) Mobley et al. (2000) observed 21 groups. Baird et al. (2008) encountered false killer whales 
in depths between 48 m and 4,331 m depth. During the 2002 survey, Barlow et al. (2004) observed two groups 
of two and 19 individuals within the northwestern portion of the Hawaiian EEZ in August and September 2002, 
respectively (Figure 6.13). During the 2007 NWHI survey, one group of false killer whales was observed near 
Lisianski Island (NMFS, unpublished data). 




Legend 

A Pseudorca crassidens Observations 

Cetacean Survey Transects (Barlow et al., 2004) 
B Main Hawaiian Islands * 

^] Papah3naumckuakea Marine National Monument Boundary 

200 400 600 800 

Kilometers 



Figure 6.13. False killer whale observations from 2002 survey of the Hawai- 
ian Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 




sa 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Short-finned Pilot Whale (Globicephala macrorhynchus) 

The short-finned pilot whale has a large, bulbous head in which the melon protrudes beyond the mouthline. 
The beak is very short and the mouthline is noticeably upturned toward the eye. The dorsal fin is falcate, set 
forward of the midbody, broad at the base and long relative to its height. Males have distinctly broader dorsal 
fins than females. The species is relatively large with females reaching lengths of 5.5 m and weights of 1,000 
kg and males reaching lengths of 6.1 m and weights of 3,000 kg. The color pattern is generally black or dark 
brown with a light gray throat patch, saddle behind the dorsal fin and streak behind the eye. 

The short-finned pilot whale is a tropical to warm-temperate species found between 50°N and 40°S. They typi- 
cally occur in deep water. 



Short-finned pilot whales are seen 
throughout the year in the MHI (Shallen- 
berger ,1981; Mobley et al., 2000; Baird 
et al., 2003). Shallenberger (1981) not- 
ed that group sizes were often greater 
than 100 individuals and rarely smaller 
than 30 individuals. Aerial surveys con- 
ducted within 46 km of the coastline 
within the MHI (1993-1998) resulted in 
73 observed groups of pilot whales with 
an average group size of 8.4 individu- 
als (Mobley et al., 2000). A summary of 
sighting data from 2000 to 2005 resulted 
in 80 observed groups of pilot whales 
with an average of 20 individuals per 
group (Baird et al., 2005a). Data from 
May and June of 2003 demonstrated 
that the average depth of sighting loca- 
tions for 17 groups of pilot whales within 
the MHI was 1,142 m (Baird et al., 2003). 
During the 2002 survey of the Hawaiian 
EEZ, Barlow et al. (2004) observed 25 

groups of pilot whales, ranging in size from 2 to 28 individuals (Figure 6.14). Four sightings were within the 
Monument boundaries (Barlow et al., 2004). During the 2007 (March-April) NWHI survey, four groups of pilot 
whales were observed within the Monument between Pearl and Hermes Atoll and Midway Atoll, near French 
Frigate Shoals and at Gardener Pinnacles (NMFS, unpublished data). 




Legend 

a Globicephala macrortiynchus Observations 

Cetacean Survey Transects (Barlow elal., 2004) 
H Main Hawaiian Islands 

I Faoahanaurnokuakea Marine National Monument Boundary 
200 400 600 aoo 



Figure 6.14. Short-finned pilot whale observations in the Hawaiian Island 
EEZ. Sources: Barlow et al., 2004; Johnston et al., 2007; NFMS unpub- 
lished; map: K. Keller. 



Killer Whale (Orcinus orca) 

The killer whale has a stocky body with a conical shaped head that lacks a prominent beak. This is the largest 
of the delphinid species with females growing to a maximum length of 8.0 m and an average weight of 3,000 
kg and males growing to a maximum length of 9.0 m and an average weight of 6,000 kg. The dorsal fin is large, 
ranging in size from 0.9 m for females to 1.8 m for males. The shape of the dorsal fin is variable, from falcate in 
females and juveniles to tall and erect in males. The color pattern of the killer whale is its most distinctive fea- 
ture. The dorsal side of the body is black, as are the pectoral flippers. The ventral side of the body is white and 
lobes extend from the belly along each side behind the dorsal fin. Distinct white patches are located slightly 
above and behind the eyes. A gray to white shaded saddle is located on the back behind the dorsal fin. 



Killer whales are considered the most widespread cetacean. They can be found in any marine region but 
are more abundant in cool temperate regions. In the Pacific Northwest, two different groups of killer whales 
have been identified. Both the "transients" and "residents" are present year-round, however the "transients" 
have larger home ranges and prey on marine mammals, while resident pods target fish as their primary prey. 
Whether this pattern is universal is unknown. Movements of killer whales appear to be driven by food avail- 
ability (American Cetacean Society, 1995 -2007). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Although infrequent, sightings of killer 
whales have been documented in the 
Hawaiian EEZ (Shallenberger, 1981; 
Tomich, 1986; Baird etal., 2005b). Baird 
et al. (2005b) reported 21 records of 
killer whale sightings within the Hawai- 
ian EEZ between 1994 and 2004. Dur- 
ing an aerial survey in March of 2000, 
Mobley et al. (2001) observed a group 
of killer whales west of Niihau. In May 
of 2003, Baird et al. (2003) observed 
one group of four individuals off the 
west side of the island of Hawaii in 773 
m depth. Barlow et al. (2004) observed 
two groups of killer whales, each con- 
sisting of two individuals, in September 
and November of 2002 (Figure 6.15). 
The November sighting occurred just 
north of French Frigate Shoals (Barlow 
etal., 2004). 




Legend 

A Orcinus orca Observations 

Cetacean Survey Transects (Bartow elal. 2004) 
Main Hawaiian islands rH 

j 1 Papahanaumokuakea Marine National Monument Boundary 

20O 400 600 aoo 

Kilometers 



Figure 6.15. Killer whale observations from 2002 survey of the Hawaiian 
Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 



Sperm Whale (Physeter macrocephalus) 

The sperm whale is large and distinctive, with a huge square head comprising approximately one-third of its 
total length. Adult females can reach 12 m in length and 24,000 kg in weight. Adult males are larger, reaching 
18 m and weighing up to 57,000 kg. The blowhole is set at the front of the head and skewed to the left. The 
dorsal fin is small and rounded. The skin behind the head is wrinkled. The color pattern is black to brownish- 
gray with white around the mouth and belly. 



Sperm whales are cosmopolitan in their distribution and inhabit waters from the equator to the edge of the polar 
ice packs. Only adult male sperm whales will move between higher and lower latitudes, while females, calves 
and juveniles are generally found in more topical and subtropical waters year around. The breeding grounds 
are located in the tropical and subtropical waters of the lower latitudes. Sperm whales are primarily found in 
waters greater than 600 m deep. 



Data from the Townsend (1935) charts 
indicate that occurrences of sperm 
whales around the Hawaiian archi- 
pelago were common (Figure 6.16). 
The whaling industry decimated their 
numbers, and currently the species is 
listed as Endangered under the ESA 
(http://www.nmfs.noaa.gov/pr/species/ 
esa/mammals.htm). Shallenberger 

(1981) reported occasional sightings 
and stranding events within the MHI. 
Sperm whales were the most frequently 
observed species within the Hawaiian 
EEZ between August and December 
2002 (Barlow et al., 2004). Barlow et al. 
(2004) observed 46 groups ranging in 
size from one to six individuals. Twelve 
of these sightings were within the Mon- 
ument boundaries (Barlow et al., 2004). 





Legend 

Physeter macrocephalus Observations 

Cetacean Survey Transects (Barlow at al., 20O4) 
H Main Hawaiian Islands 
I Papahanaumokuakea Marine National Monument Boundary 

200 400 800 800 

u Kilometers 



i 




Figure 6.16. Sperm whale observations around the Hawaiian archipelago 
Sources: Barlow et al., 2004; Johnston et al., 2007; NFMS, unpublished; 
Townsend, 1935; map: K. Keller. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

During the 2007 (March-April) NWHI survey, seven groups of sperm whales were observed within the Monu- 
ment (NMFS, unpublished data). 



Pygmy Sperm Whale (Kogia breviceps) 

The pygmy sperm whale has a small, robust body with a bulbous, square-shaped head with a pointed snout, 
a narrow mouth and undershot lower jaw. It ranges from 2.8 m to 3.5 m in length and reaches a maximum 
weight of 450 kg. The blowhole is set far back on the head and offset to the left. The dorsal fin is small, falcate 
and located behind the midpoint of the body. The pectoral flippers are small and located unusually close to the 
head. Coloration ranges from a dark gray on the back to pinkish-white on the belly, and there are half-moon 
shaped markings between the eye and the flipper, sometimes referred to as "false-gills". The pygmy sperm 
whale closely resembles the dwarf sperm whale. 

The pygmy sperm whale occurs in tropical to temperate regions worldwide and is generally found in deep wa- 
ters seaward of the continental shelf. Pygmy sperm whales are thought to be non-migratory and are generally 
found singly or in small groups up to five individuals. 

Pygmy sperm whales are observed 
occasionally within Hawaiian waters. 
Shallenberger (1981) combines the two 
Kogia species into one section because 
of the possible misidentification of each 
species. Most data come from stand- 
ings in the MHI, which have occurred on 
Kauai, Oahu, Molokai and Maui (Shal- 
lenberger, 1981). In June 2003, Baird 
et al. (2003) observed one group of two 
individuals between Kauai and Niihau 
in 700 m depth. Two lone individuals 
were observed within the northwestern 
portion of the Hawaiian EEZ by Barlow 
et al. (2004; Figure 6.17) in November 
2002. Within the Monument there is a 
single record, from 1923, of a beach 
cast skeleton that was identified as a 
pygmy sperm whale (Amerson, 1971). It 
was this event that led to the naming of 
Whale Island at French Frigate Shoals 
(Amerson, 1971). 




Legend 

A Kogia bfeviceps Observations 

Cetacean Survey Transects (Bartow et al., 2004) 
Main Hawaiian Islands i\ 

| Papahanaumokualwa Marina National Monument Boundary 
200 



Figure 6.17. Pygmy sperm whale observations from 2002 survey of the 
Hawaiian Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 



Dwarf Sperm Whale (Kogia sima) 

The dwarf sperm whale is nearly identical in appearance to the pygmy sperm whale; however, the dwarf sperm 
whale has a dorsal fin that is more prominent and positioned further forward on its back. Additionally, the dwarf 
sperm whale is a smaller whale, reaching a length of 2.7 m and weighing approximately 275 kg. 

The dwarf sperm whale is a tropical and temperate species, generally found in offshore waters, but somewhat 
more coastal than the pygmy sperm whale. There is no evidence to suggest that the dwarf sperm whale mi- 
grates. 

Shallenberger (1981) combines the two Kogia species into one section because of the possible misidentifica- 
tion of each species. Stranding events have been recorded on Oahu and Hawaii (Tomich, 1986). In May and 
June of 2003, Baird et al. (2003) encountered eight groups of dwarf sperm whales near Lanai, Kauai and Nii- 
hau. The average group size was two individuals, and they were observed in waters with an average depth of 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Legend 

A Kogia sima Observations 

Cetacean Survey Transects (Barlow et al.. 2004} 
H Main Hawaiian islands 
I Papahanaumokuakea Marine National Monument Boundary 

200 400 600 SOO 

n Kilometers 



Figure 6.18. Dwarf sperm whale observations from 2002 survey of the Ha- 
waiian Island EEZ. Source: Barlow et ai, 2004; map: K. Keller. 



2,004 m (Baird et al., 2003). Barlow et al. 
(2004) observed dwarf sperm whales on 
five occasions within the Hawaiian EEZ 
in October and November 2002 (Figure 
6.18). The species was observed singly 
or in groups of up to three individuals 
(Barlow et al., 2004). 



Blainville's Beaked Whale 
(Mesoplodon densirostris) 
The Blainville's beaked whale, some- 
times called the dense-beaked whale, 
has a robust body that is compressed 
laterally. It is a medium-sized whale that 
reaches 5.8 m in length and 3,500 kg in 
weight. The dorsal fin is small, falcate 
and positioned approximately two-thirds 
down the length of the body. The color 
pattern is silver-gray to brown on the 
dorsal surface with lighter gray to white 

on the belly. Blainville's beaked whales are typically covered with white or pale-gray circular marks or scars 
that are likely caused by cookie cutter sharks. They have a distinct mouthline that sweeps up at the middle of 
the lower jaw. Males have a tooth protruding from each side of the lower jaw at this site. 

Blainville's beaked whales are a tropical and temperate species and are considered the most widely distributed 
mesoplodont. They are found primarily in deep waters, 500-1,000 m. There is no evidence of seasonal move- 
ments or migrations. 

Blainville's beaked whales have been 
observed in the MHI off of Oahu, Ha- 
waii, Kauai and Molokai (Shallenberg- 
er, 1981; Tomich, 1986; Mobley et al., 
2000, Baird et al., 2003). Group sizes 
range from two to seven individuals 
(Shallenberger, 1981; Tomich, 1986; 
Mobley et al., 2000; Baird et al., 2003). 
Five groups observed by Baird et al. 
(2003) were in average water depths of 
1,304 m. Recent satellite tagging stud- 
ies off the west coast of the island of 
Hawaii suggest that this population of 
Blainville's beaked whales is island-as- 
sociated and exhibits strong site fidelity 
(Schorr et al., In press). Within the Mon- 
ument boundaries, the earliest record 
of Blainville's beaked whales is from 
a stranding event of two individuals at 
Midway Atoll in April 1961 (Galbreath, 
1963; Shallenberger, 1981). Barlow et 
al. (2004) observed Blainville's beaked whales (one to two individuals per sighting) within Monument waters in 
September and October 2002 (Figure 6.19). 




Legend 

A Mesoototfon densitoslrts Observations 

Ceiacean Survey Transects (Bartow el al.. 2004) 
| Main Hawaiian Islands f , 

I Papahanaumofcuaivse Marine National Monument Boundary 

BOO 
3 Kilometers 



Figure 6.19. Blainville's beaked whale observations from 2002 survey of 
the Hawaiian Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 




W 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Cuvier's Beaked Whale (Ziphius cavirosths) 

Cuvier's beaked whale has a long, stocky body. Adults range in length from 5 m to 7.5 m and weigh from 2,000 
kg to 3,000 kg. The head is small, the melon is steeply tapered and the beak is short and poorly-defined. Ma- 
ture males have two teeth protruding from the front of the lower jaw. The dorsal fin is small, falcate and located 
approximately two-thirds back along the length of the body. The color pattern varies from dark gray to light 
brown, with the head and neck, and eventually the body, becoming lighter in color as the whale ages. This is 
particularly pronounced in males. Scars from cookie cutter sharks can give a mottled appearance to the sides 
and belly. 

Cuvier's beaked whales occur in all offshore waters except those in the polar regions. They are primarily found 
in waters greater than 1,000 m deep. 



Cuvier's beaked whales are observed 
infrequently within the Hawaiian EEZ. 
Shallenberger (1981) noted two sight- 
ings off of Lanai and Maui. Aerial surveys 
of the MHI conducted during 1993, 1995 
and 1998 resulted in seven observed 
groups of Cuvier's beaked whales with 
an average group size of three individu- 
als (Mobley et al., 2000). During a 10 
year period (1990-2006), McSweeney 
et al. (2007) encountered Cuvier's 
beaked whales on 35 occasions off the 
west coast of the island of Hawaii. Re- 
sightings of individuals during this time 
suggest some degree of site fidelity and 
the presence of a resident population. 
Stranding events at Midway Atoll and 
Pearl and Hermes Atoll demonstrate 
the presence of Cuvier's beaked whales 
within the Monument (Shallengberger, 
1981). In addition, Barlow et al. (2004) 
observed groups within the Monument, 
at Gardener Pinnacles and north of Mo- 
kumanamana in August of 2002 (Figure 
6.20). 



Longman's Beaked Whale 
(Indopacetus pacificus) 
The Longman's beaked whale, some- 
times referred to as the tropical bottle- 
nose whale, can reach 9 m in length. 
It has a bulbous head and moderately 
long beak, from which two teeth erupt 
in males. The dorsal fin is pointed, fal- 
cate, and located behind the midpoint of 
the body. The color varies from brown to 
bluish gray, and the head and sides can 
be a lighter color in younger animals. 
This species has been misidentified as 
the southern bottlenose whale. 




Figure 6.20. Cuvier's beaked whale observations from 2002 survey of the 
Hawaiian Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 




Legend 

a Indopacetus pacificus Ob&arvadons 

Cetacean Survey Transects (Barlow et al.. 2004) 
H Main Hawaiian Islands 
1 Papahanaurnokuakaa Marine National Monument Boundary 



Figure 6.21. Longman's beaked whale observations from 2002 survey of 
the Hawaiian Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Little is known about the range of Longman's beaked whale, but it is thought to occur primarily in pelagic waters 
within the Indo-Pacific region. 

Longman's beaked whale is rare in Hawaiian waters. There are only two confirmed sightings within the Hawai- 
ian EEZ. The first was made in November 2002 when Barlow et al. (2004) observed one group of four individu- 
als within the northwestern portion of the EEZ (Figure 6.21). The second confirmed sighting was a group of 
30-35 individuals off of the west coast of the island of Hawaii in 2007 (http://www.cascadiaresearch.org/robin/ 
August2007.htm). 

Minke Whale (Balaenoptera acutorostrata) 

Minke whales are the smallest of the baleen whales. At least two subspecies of the common minke whale are 
recognized and include the North Pacific (B.a.scammoni) and North Atlantic (B.a. acutorostrata). The dwarf 
minke whale is considered a potential third subspecies but has not been given an official scientific name. The 
common minke whale has a small, slender body and a pointed triangular head with a well-defined longitudinal 
ridge along the rostrum. Adults range in length from 7 m to 10.7 m and weigh as much as 9,200 kg. Females 
are slightly longer than males. The falcate dorsal fin is set approximately two-thirds back along the length of 
the body. The color pattern is black or dark gray on the dorsal side with a lighter gray chevron across the back 
and white on the belly. A white band across the middle of the pectoral flippers is a distinctive characteristic of 
this species. 

Common minke whales occur in the 
North Atlantic and North Pacific and mi- 
grate from the polar and temperate wa- 
ters in the summer to the tropical waters 
in the winter. They are frequently ob- 
served in coastal or shelf waters, rath- 
er than deep offshore habitats. In the 
eastern Pacific, minke whales are found 
from the Bering Sea south to the coast 
of Baja California, and in the western 
Pacific they are found from the Sea of 
Okhotsk to the Sea of Japan. The win- 
ter distribution of North Pacific minke 
whales can be inferred from the distri- 
bution of their distinctive calls (termed 
"boings"). Boings are heard primarily be- 
tween 15°N and 30°N in the months of 
November through March (Rankin and 
Barlow 2005). Shallenberger (1981) lists 
the minke whale under "animals sighted 
in Hawaiian waters but not considered 
part of the normal cetacean fauna." One 
minke whale was observed within the Hawaiian EEZ by Barlow et al. (2004) in November 2002 (Figure 6.22). 

Bryde's Whale (Balaenoptera edeni) 

The Bryde's whale has a long, slender body with a pointed rostrum. Females are larger than males and can 
reach a length of 15.5 m and a weight of 40,000 kg. Three ridges extend from the blowhole to the tip of the 
rostrum, which is a diagnostic feature of this species. The dorsal fin is extremely falcate, tall (up to 45 cm), and 
positioned two-thirds of the way along the length of the body. The color pattern is gray on the dorsal side and 
white on the belly, sometimes with banding or chevrons. 




Figure 6.22. Minke whale observations from 2002 survey of the Hawaiian 
Island EEZ. Source: Barlow et al., 2004; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Bryde's whales are found in tropical and subtropical waters of the Pacific, Atlantic and Indian Oceans. They are 
rarely seen north or south of 40°. The whales are often seen at nearshore upwellings but are also occasion- 
ally found offshore. Some evidence suggests that there are two forms of the Bryde's whale, an inshore and an 
offshore that may differ in reproductive cycles and diet. The Bryde's whale is not as migratory as the sei whale, 
but limited migration does take place in some populations (offshore form), while other populations are resident 
year-around (inshore form). 



Shallenberger (1981) listed the Bryde's 
whale as a rare species in Hawaiian 
waters. He noted one confirmed sight- 
ing 87 km southeast of Nihoa in April 
of 1977 (Shallenberger, 1981). Tomich 
(1986) cites Leatherwood et al. (1982) 
as noting that Bryde's whales are "rela- 
tively abundant over shallows northwest 
of Hawaii and near Midway Islands." 
During the 2007 NWHI survey, a single 
Bryde's whale was observed off the east 
coast of Niihau (NMFS, unpublished 
data). During August through October 
of the 2002 Hawaiian EEZ survey, Bar- 
low et al. (2004) observed 14 groups of 
Bryde's whales (Figure 6.23). With the 
exception of two sightings, in which 
there were two whales present, all were 
of single whales. Six of the sightings 
were within the Monument boundaries 
(Barlow etal., 2004). 




Legend 

A Bateerjapte/a etfew Observations 

Cetacean Survey Transects (Bartow et el.. 2QTJ4) 
Mam Hawaiian Islands 
I Papahanaumokuakea Marine National Monument Boundary 
u 200 400 eon sou 

3 Kilometers 



Figure 6.23. Byrde's whale observations in the Hawaiian Island EEZ. 
Sources: Barlow et al., 2004; Johnston et al., 2007; NMFS, unpublished; 
map: K. Keller. 



Sei Whale (Balaenoptera borealis) 

Sei whales are similar in appearance to Bryde's whales, with a gray dorsal and white ventral surface, and with 
the dorsal fin rising at a steep angle from 
the back. Adults reach a maximum of 
20 m in length, and may weigh as much 
as 45,000 kg. Sei whales have only a 
single rostral ridge and lack the two ad- 
ditional parallel longitudinal ridges that 
are evident in the Bryde's whales. 

Sei whales are found from the tropics to 
polar regions in the northern and south- 
ern hemispheres, but most often occur 
in the mid-latitude temperate zones. 
They are migratory open-ocean whales, 
not often observed near the coasts. The 
species is listed as Endangered under 
the ESA (http://www.nmfs.noaa.gov/pr/ 
species/esa/mammals.htm). 

Neither Shallenberger (1981) nor To- 
mich (1986) mention sei whales. Indi- 
vidual sei whales were observed on 
several occasions within the Hawaiian 




Legend 

a Balaenoptera borealis Observations 

Cetacean Survey Transects (Barlow etal.. 
H Main Hawaiian Islands 

I Papahanaumokuakea Marine National Monument Boundary 
200 400 600 000 Kilometers 



Figure 6.24. Sei whale observations in the Hawaiian Island EEZ. Sources: 
Barlow et al., 2004; Johnston et al., 2007; NMFS, unpublished; map: K. 
Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

EEZ (but outside of Monument waters) by Barlow et al. (2004) in November 2002 (Figure 6.24). In April 2007, 
during the NWHI survey a single sei whale was observed between Laysan and Lisianski Islands (NMFS, un- 
published data). 

Fin Whale (Balaenoptera physalus) 

Similar to the Bryde's and sei whales, fin whales are dark brown or gray dorsally and white ventrally. Fin whales 
are significantly larger reaching a length of 24 m and a weight of 120,000 kg. The dorsal fin is variable and ei- 
ther pointed or falcate. Fin whales have a distinctive asymmetrical color pattern on the lower jaw; the right side 
(including baleen) is white and the left side is dark gray or black. In addition, most individuals have a v-shaped 
chevron across the back of the head and a swirled blaze on the right side of the head. 



Fin whales are found in all oceans of the world, but primarily occur in cooler temperate regions and concentrate 
on shelf and in coastal waters. They can be found over a broad latitudinal range throughout the year, however, 
some appear to migrate, spending the summer in the northern polar region and the winter in warmer waters of 
lower latitudes. Like the sei whale, fin whales are listed as Endangered under the ESA (http://www.nmfs.noaa. 
gov/pr/species/esa/mammals.htm). 



Shallenberger (1981) listed fin whales 
as rare in Hawaiian waters. He report- 
ed on two sightings off of Oahu and 
one stranding on Maui (Shallenberger, 
1981). In February 1994, a single fin 
whale was observed near Kauai (Mob- 
ley et al. 1996). Acoustic recordings 
have also detected the presence of fin 
whales off Oahu and Midway (Thomp- 
son and Friedl, 1982; McDonald and 
Fox, 1999). During the period of Sep- 
tember through November 2002, Bar- 
low et al. (2004) observed five groups 
of fin whales within the Hawaiian EEZ 
(Figure 6.25). A single individual was 
observed within the Monument between 
Maro Reef and Laysan Island (Barlow 
etal.,2004). 




Legend 

A Balaenoptera physalus Observations 

Cetacean Survey Transects (Barlow elal.. 2004) 
H Main Hawaiian Islands 
I I PapahSnaumokuSkea Marine National Monument Boundary 



Figure 6.25. Fin whale observations from 2002 survey of the Hawaiian Is- 
land EEZ. Source: Barlow et al., 2004; map: K. Keller. 



Humpback Whale (Megaptera novaeangliae) 

The humpback whale has a large, robust body that reaches a maximum length of 17 m and a maximum weight 
of 40,000 kg. Females are slightly larger than males. Humbacks have very long flippers (up to one-third the 
length of the body). The head and lower jaw are covered in tubercles. The dorsal fin is variable from tall and 
falcate to a small hump. The large flukes are concave with a serrated trailing edge. The color pattern is vari- 
able. It can be black on the dorsal side and black, white, or mottled black and white on the ventral side. 

Humpback whales are found in all of the major oceans and occur primarily in coastal waters (Figure 6.26). 
They are a migratory species and spend the summer in the polar regions and the winter in tropical waters. 
Four stocks are believed to occur in the North Pacific, including one that winters in the central North Pacific 
and Hawaii. Multiple stocks also occur in the North Atlantic, Northern Indian Ocean and in the southern hemi- 
sphere. Humpback whales are listed as Endangered under the ESA (http://www.nmfs.noaa.gov/pr/species/ 
esa/mammals.htm). 



Humpback whales within the MHI have been extensively studied for nearly four decades (Herman andAntinoja, 
1977; Shallenberger, 1977; Herman et al., 1980; Shallenberger, 1981; Calalmonkidis et al., 2008) and individ- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




± Humpback whale 

V Probable humpback whale 

',..; Acoustic detections 
^B -6,100 to -5,000 m 
(■I -5,900 to -5,000 m 
^■-4,900 to -4,000 m 
■ -3,900 to -3,000 m 

— |-Z,900to-Z,000m 

31 - 1 , 900 to -1,600 m 

— 1-1,400 to -1,000 m 
I 1 -900 to -500 m 
-490 to -200 m 
■■ > -200m 

Monument boundary 

— — 21.1 C Synoptic 

21 .TCJan -March 200T 

^— On effort 






<A 



9, 






Nihoa 

Necker Island (Mokurnanamana) 
: Gardner Pinnacles 

Mara Reef 

Laysan Island 

Lisianski Island 
- Pearl & Hermes Reef 

Midway Atoll 

Ladd Sea mount 



o 






T 



■J 



Figure 6.26. Humpback whale modeled habitat. A = Nihoa; B = Mokurnanamana; C = Gardner Pinnacles; D = Maro Reef; 
E = Laysan Island; F = Lisianski Island; G = Pearl and Hermes Atoll; H = Midway Atoll; and I = Ladd Seamount. Map: 
Johnston et al., 2007. 



ual whales photographed in Hawaii are 
also known from records in the northern 
feeding grounds in Alaska and British 
Columbia (Calambokidis et al., 2008). 
The whales migrate to the waters sur- 
rounding the Hawaiian Islands begin- 
ning in November and remain there un- 
til late May (Shallenberger, 1981). While 
in Hawaiian waters, they are found ex- 
clusively in shallow waters less than 
183 m (Shallengberger, 1981; Johnston 
et al., 2007). Humpback whales may 
use the waters of the NWHI throughout 
the winter months, and were observed 
within the Monument boundaries near 
Nihoa, Mokurnanamana, Gardner Pin- 
nacles, Maro Reef and Lisianski Island 
(Johnston et al., 2007). Nine groups 
were observed, two of which contained 
calves (Johnston et al., 2007; Figure 
6.27). Johnston et al. (2007) modeled 
the availability of humpback whale win- 
tering habitat within the Monument based on bathymetric and sea surface temperature data. This modeling in- 
dicates potential available humpback whale habitat at all islands and atolls in the NWHI (and at Maro Reef and 
Ladd Seamount) south of the 21.1°C sea surface temperature dine. These areas in 2007 totaled 14,700 km 2 , 
compared to 7,200 km 2 of humpback wintering habitat available in the MHI (Johnston et al., 2007). Surveys 
(aerial and ship-based) of the NWHI conducted in 1976-1977 (November-April) returned no sightings of hump- 




Legend 

A Megaptera novaeangtiae Observations 

Cetacean Survey Transects (Bartow et al., 2004) 
5$ Main Hawaiian Islands 
1 ^J Papahanaumokuakea Marine National Monument Boundary 

a :cs <» sou im 



Figure 6.27. Humpback whale observations in the Hawaiian Island EEZ. 
Sources: Barlow et al., 2004; Johnston et al., 2007; NMFS, unpublished; 
map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

back whales (Herman et al., 1980). However, since the mid-1990s, regular sightings of humpback whales each 
year from January through March have been made by U.S. Fish and Wildlife Service staff stationed at French 
Frigate Shoals (A. Anders, pers. comm.). One reported opportunistic sighting of a mother and calf at French 
Frigate Shoals in February 1977 was dismissed as a "straggler" (Herman et al., 1980). This may suggest that 
the NWHI provide winter habitat for an expanding humpback whale population (Johnston et al., 2007). 

Additional Cetacean Species 

In addition to the 23 species described above, two other cetacean species have been detected within the Ha- 
waiian Islands EEZ, including the North Pacific right whale (Eubalaenajaponica) and blue whale (Balaenoptera 
musculus; Barlow, 2006). North Pacific Right whales have been observed in the waters surrounding the MHI 
on two confirmed occasions (Rountree et al.,1980;Tomich, 1986). The first record comes from a whaling ship's 
logbook from 1851 in which a single "straggler" was observed 250 nm west of Maui (Rountree et al., 1980). 
Tomich (1986) noted a sighting of an individual right whale swimming among a group of humpback whales 
between Maui and Lanai in March 1979. Blue whales have been detected, based on vocalizations, off of the 
coast of Oahu and it was suggested that they migrate past the Hawaiian Islands twice a year (Thompson and 
Friedl, 1982). Twenty hertz signals, similar to those recorded by Thompson and Friedl (1982), have also been 
reported near Midway Atoll (Tomich, 1986). 

Cetacean Abundance Estimates 

The 2002 cetacean survey conducted by Barlow et al. (2004) in the Hawaiian Islands EEZ allowed for the cal- 
culation of abundance and density estimates for 19 of the 23 species observed (see Barlow 2006 for descrip- 
tion of analytical methods). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



PINNIPEDS 

The Hawaiian monk seal (Monachus 
schauinslandi; Figure 6.28) was listed 
as Depleted under the MMPA and En- 
dangered under the ESA in 1976. The 
current population estimates are around 
1,100-1,200 individual seals, the major- 
ity of which live in the NWHI. 

The genus Monachus is a wide ranging 
species and is found in several differ- 
ent geographic areas around the world 
(Figure 6.29). The genus includes the 
Mediterranean monk seal (Monachus 
monachus), the Caribbean monk seal 
(Monachus tropicalis) and the Hawaiian 
monk seal. The Mediterranean monk 
seal is critically endangered and the 
Caribbean monk seal is assumed to 
have gone extinct in the last 50 years 
(Kenyan, 1977; see Boyd and Stan- 
field). The Hawaiian monk seal popula- 
tions are estimated to have declined by 
60% since the 1950s (Antonelis et al., 
2006). 



Range 

Based on anatomical features and DNA 
analysis researchers estimate that Ha- 
waiian monk seals arrived in Hawaii 
14-15 million years ago (Repenning et 
al., 1979) and split from the Monachus 
ancestors around 11.8 to 13.8 million 
years ago (Flyer et al., 2005). Hawaiian 
monk seals occur within the Hawaiian 
EEZ and the main subpopulations oc- 
cur in the NWHI. A smaller but poten- 
tially increasing population of seals 
inhabit the MHI and there have been 
occurrences including a documented 
pupping and relocations of aggressive 
males to Johnston Atoll. The monk seal 
metapopulation can be divided into six 
major and two smaller subpopulations 
in the NWHI and one in the MHI (Figure 
6.30). These subpopulations are further 
grouped into management units. 





Figure 6.28. There are an estimated 1,200 Hawaiian monk seals, the ma- 
jority of which live in the NWHI. In 1976 the species was listed as Endan- 
gered under the ESA. Photo: J. Watt. 



j* -.5? 




Legend 
Monk Seals 

A Carribean Monk Seal, Extinct 
■ Hawaiian Monk Seals, Endangered 
# Mediterranean Monk Seal, Critically Endangered 
I — | Papahanaumokuaeakea Marine National Monument 
Continents 



2.000 4,000 6,000 



B.aoo 

^ Kilometers 



Figure 6.29. Worldwide distribution of monk seais. Source: ESRI; map: K. 
Keiier. 




Legend 

Monk Seal Subpopulations 

Western Subpopulations 
■ Isolated Sub populations 
* Limited Data 
J | | Papahanaumokuakea Marina National Monument Boundary 



100 200 300 400 

i Kilometers 



Figure 6.30. Hawaiian monk seal subpopulations. Source: NMFS, unpub 
data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Available Data 

The U.S. Fish and Wildlife Service conducted periodic monitoring of Hawaiian monk seals from the late 1950s 
until the late 1970s when the National Marine Fisheries Service (NMFS) assumed responsibility for recovery of 
the species and commenced monitoring activities. NMFS began an annual monitoring of Hawaiian monk seals 
at most major sites in the early 1980s. There is no historic data for estimating population size prior to the sur- 
veys of the 1950s or to estimate carrying capacity. It is likely that upon arriving in the MHI, Polynesian settlers 
extirpated the local population of seals. The surveys of the NWHI in the 1950s were too soon after World War 
II for the population to have plausibly recovered from the impact of the military disturbance, and some military 
presence was still having a negative effect on the monk seals at that time. 

The first range-wide surveys of Hawai- 
ian monk seals were conducted in the 
late 1950s (Kenyon and Rice, 1959; 
Rice, 1960). Additional counts were 
conducted at Midway Atoll in 1956- 
1958 (Rice, 1960) and at Kure Atoll in 
1963-1965 (Wirtz, 1968). Surveys were 
repeated throughout the 1960s and 
1970s, and while the methods were not 
standardized, complete beach counts 
are roughly comparable between the 
two survey periods. 

Since the early 1980s NMFS has been 
conducting annual surveys using stan- 
dardized methods to estimate the popu- 
lation of Hawaiian monk seals. The work 
is conducted at field camps that are ac- 
tive from approximately April through 
August (Figure 6.31). All of the field 
camps collect information on the num- 
bers of seals in the subpopulations based on counts of uniquely-identified seals. In addition any births, deaths, 
serious injuries and entanglements are documented. Necropsies are conducted on dead seals and seal feces 
are collected to evaluate dietary information. 

Habitat Use and Foraging Behavior 

Between 1996 and 2002, the movements and diving patterns of 147 Hawaiian monk seals have been moni- 
tored with satellite-linked radio transmitters at the six breeding colonies in the NWHI (42 adult males, 35 adult 
females, 29 juvenile males, 14 juvenile females, 12 weaned male pups, 15 weaned female pups; Abernathy 
and Siniff,1998; Stewart, 2004a, b; Stewart and Yochem, 2004a, b, c). 

Parrish et al. (2000) attached animal borne imaging systems (Crittercams) to 24 adult and subadult male monk 
seals at French Frigate Shoals. The Crittercams recorded the habitat depth and bottom type at locations where 
monk seals were seen capturing prey items. Recent studies have focused on characterizing juvenile monk 
seal habitat use and foraging behavior at French Frigate Shoals using Crittercams and time-depth recorders 
(TDRs). 




Figure 6.31. Hawaiian monk seal field camps. Map: K. Keller. 



Abundance 

NMFS field camps were initiated in the early 1980s using systematic surveys for estimating abundance of 
the Hawaiian monk seal populations. The abundance of monk seals at the six main reproductive sites in the 
NWHI is estimated by direct enumeration. At those locations the majority of individual seals can be identified by 
fiipper-tags that have routinely been applied to weaned pups since the early 1980s, bleach marks placed annu- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

ally, and by natural features such as scars and distinctive pelage patterns (Harting et al., 2007). The methods 
are different at Mokumanamana and Nihoa Islands because they are difficult to reach and there are no regular 
field camps at those sites. 

The methods used to derive estimates of monk seal abundance, and the abundance at locations within the 
Monument and in the MHI are as follows: 

• Main reproductive sub-populations in the NWHI of French Frigate Shoals, Laysan, Lisianski, Pearl and 
Hermes Atoll, Midway Islands and Kure Atoll: total enumeration of individuals when possible, otherwise 
capture-recapture estimates or minimum abundance (Baker, 2004; Baker et al., 2006). 

• Mokumanamana and Nihoa Islands: corrected mean beach counts made during most recent five years at 
Mokumanamana and Nihoa Islands. A correction factor (2.89 ± 0.06; NMFS, unpubl. data) derived from 
observations at the main reproductive sites is applied. 

• MHI: Minimum abundance consisting of the total number of uniquely identifiable seals observed alive dur- 
ing a calendar year. Sightings are non-systematic and collected by NMFS, and reported by volunteers, 
partner agencies and the general public. 

Beach Counts 

Methods 

Beach counts are conducted at least eight times annually per site to calculate a mean value that serves as a 

trend index for long-term comparisons. The beach counts include a count of all the seals found on the island 

or group of islands within an atoll during a single mid-day survey. 

Uses 

Direct enumeration data cannot be used for comparing historical counts; however, a measure of long-term 
trend is derived from the mean of all of the beach counts that have been conducted with varying frequency 
since the late 1950s. These beach counts provide a useful index because the general methodologies of counts 
during these 45 years are roughly comparable. 

Limitations 

A consideration when interpreting the mean beach counts is that the relationship between the mean beach 
counts and the actual population size is uncertain. That is, all of the factors that might cause beach counts to 
deviate from the true abundance (for example, changes in haul out patterns over time) are not known, and 
hence appropriate correction factors have not been determined. Eberhardt et al. (1999) concluded that, "beach 
counts may be very poor guides to year-to-year trends. However, beach counts are valuable indicators of 
long-term trends." NMFS is currently investigating other approaches for estimating total abundance to better 
characterize long- and short-term trends. 

Habitat Use 
Terrestrial Habitat Use 

Monk seals use terrestrial habitat for haul-out areas to rest and pupping. Haul-out areas for resting generally 
consist of sandy beaches, but virtually all substrates, including emergent reef and shipwrecks, are used at vari- 
ous islands. Monk seals also use the vegetation behind the beaches, when available, as a shelter from wind 
and rain. Pups are born on various substrates; however, sandy beaches with shallow protected water near 
shore seem to be preferred habitat for pupping and nursing (Westlake and Gilmartin, 1990). 

Marine Habitat Use 

Monk seals spend approximately two-thirds of their time in the marine habitat (MMRP, unpublished data). They 
are primarily benthic foragers (Goodman-Lowe et al., 1998), and will search for food in coral reef habitat and 
on substrate composed of talus and sand on marine terraces of atolls and banks to depths exceeding 500 
m (Parrish et al., 2000, 2002: Parrish and Abernathy, 2006). Parrish et al. (2002) also described monk seals 
foraging in corals below 300 m in subphotic zones (Parrish et al., 2002). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Legend 

Foraging Ranges 
I I Papahanaumokuakea Marine National Monument Boundary 



The largest study of monk seal foraging 
ranges and diving behavior was con- 
ducted using tagging information from 
1996 to 2002 (Abernathy and Siniff, 
1998; Stewart 2004a, b; Stewart and 
Yochem, 2004a, b, c). During this time 
the movements and diving patterns of 
147 Hawaiian monk seals were moni- 
tored with satellite-linked radio trans- 
mitters at the six breeding colonies in 
the NWHI. Spatial dispersal of forag- 
ing seals indicates that they forage ex- 
tensively within the lagoons at French 
Frigate Shoals, Pearl and Hermes Atoll, 
Midway Atoll and Kure Atoll, and on the 
outer slopes of those atolls and sea- 
ward of Laysan and Lisianski Island. 
Seals also foraged along the submarine 
ridges between these atolls and islands 
and at virtually all nearby seamounts. 
A fixed kernel density estimate method 
was used to determine the extent of foraging areas (Stewart et al., 2006; Figure 6.32). Primary foraging oc- 
curred in areas with high bathymetric relief or focused within the lagoon areas. At the majority of the islands 
and atolls 95% of the foraging occurred within 38 km and 75% occurred within 20 km of center of the island or 
atoll (Stewart et al., 2006). French Frigate Shoals did not follow this same pattern, however, with 95% of the 
foraging occurring within 50-58 km of the center of the atoll (Stewart et al., 2006). Seals at all of the colonies 
foraged outside of the colonies but there was no distinct pattern (Stewart et al., 2006; Table 6.2). Distances 
traveled to forage from haul out sites also varied with a seal's age and sex and with the seal's colony of seal 
origin. Foraging distances ranged overall from less than 1 km up to 217 km (Abernathy, 1999; Stewart, 2004a, 
b; Stewart and Yochem, 2004a, b, c; Stewart et al., 2006) 




400 
DKilometers 



Figure 6.32. Hawaiian monk seal foraging ranges. Source: Stewart et al., 
2006; map: K. Keller. 



In addition to looking at the distances seals foraged from their colonies, the diving behavior was also moni- 
tored (Stewart et al., 2006; Table 6.2). Most frequently, seals dove to depths less than 150 m, though there 
were secondary diving modes at various depths up to 500 m. There was some variation in seals foraging 
depth between the island and atolls. At 
Pearl and Hermes Atoll 90 % of dives 
and at French Frigate Shoals 60 - 80% 
of dives were to depths of less than 40 
m. The remaining 10 - 20% of dives oc- 
curred in depths greater than 40 m with 
some occurring as deep at 500 m. At 
Kure Atoll, Midway Atoll, Lisianski Island 
and Laysan Island seals regularly dived 
to depths greater than 40 m (Stewart et 
al., 2006). 



Foraging areas seem to vary between 
age groups, islands and individuals 
(Figure 6.33). Weaned pups at Kure 
Atoll and Midway Atoll did not range as 
far as adults whereas at Lisianski Island 
and Laysan Islands, adults and weaned 
pups exhibited similar foraging distanc- 
es (Stewart et al., 2006). Further analy- 




Legend 






Monk Seal Tracking 


Contours (meters) 


. Saal #79731 


-499- 


1D0 


Seal #79740 


— -1499 


-500 


. Seal #79746 


2499 


-1600 


Foraging Ranges -2999 


-2600 


B Emergent Land 


3000 


N 


10 20 30 


to 

a Kilometers 


A 



Figure 6.33. Examples of individual seal movements at Lisianski Island. 
Source: NMFS, unpub data; Stewart et al., 2006; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 6.2. Percentages of monitored Hawaiian monk seals at each colony 
Source: Stewart et al., 2006. 


that 


foraged at 


various sites in 


the NWHI. 


FEATURE 

NAME 


KURE 
ATOLL 


MIDWAY 
ATOLL 


PEARL AND 

HERMES 

ATOLL 


LISIANSKI 
ISLAND 


LAYSAN ISLAND 


FRENCH 
FRIGATE 
SHOALS 


WP 
(5) 


J 

(11) 


AM 
(4) 


AF 

(4) 


WP 
(4) 


J 

(8) 


AM 
(2) 


AF 
(3) 


J 
(6) 


AM 
(9) 


AF 
(9) 


WP 
(8) 


J 
(9) 


AM 
(4) 


AF 
(5) 


WP 
(10) 


j 

(10) 


AM 
(5) 


AF 
(5) 


AM 
(17) 


AF 
(17) 


Un-named 
Kure 
Seamount 1 






50 






































Un-named 
Kure 
Seamount 2 






50 






































Un-named 
Kure 
Seamount 3 






50 






































Kure Atoll 


100 


100 


100 


100 




13 


50 


33 




























Nero 
Seamount 






25 


25 




13 


50 


33 




























Midway Atoll 








25 


100 


100 


100 


66 




























Ladd 
Seamount 








25 








33 


100 


100 


100 






















Pearl and- 

Hermes 

Atoll 
















33 




























Un-named 
PHR 












































Lisianski/ 
Neva Shoals 
























100 


100 


100 


100 














Pioneer 
Bank 
























38 


22 




20 














Northamp- 
ton W 


























11 






30 


30 


20 


40 






Northamp- 
ton E 


























11 










40 


40 






Laysan 


























22 




20 


100 


90 


100 


100 






Un-named 
Laysan 1 


























11 


















Maro Reef 


























22 






70 


60 


60 


80 






Raita Bank 
































30 


30 


40 


40 






Gardner 
Pinnacles 








































6 


30 


St. Rogatien 
Banks 








































12 


10 


Brooks Bank 








































59 


30 


French Frig- 
ate Shoals 








































100 


80 


Mokumana- 
mana 










































10 



sis was conducted to examine the relationship between habitat use and diet composition. In the early 1990s 24 
seals were tracked with satellite receivers and depth recorders at French Frigate Shoals (Parrish et al., 2000). 
In this study habitat use and diet of the monk seals were compared at different ecological zones. The analysis 
indicates that the seals foraging was focused more on the transitional slope areas. Generally the seals used all 
of the ecological zones proportionally to available habitat. The study also looked at the seals' diet and the avail- 
ability of prey species in each zone. The diet derived from scat analysis did not correlate with the prey species 
composition in any of the individual zones which was expected as seals generally used multiple habitats over 
the course of foraging trips. An analysis of the dissimilarity index of prey biomass density found that the prey 
guilds documented in the bank and slope areas had the least deviation (Parrish and Abernathy, 2006). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Satellite tagging data provide basic information on where the seals travel but do not provide specific informa- 
tion about seal behavior during their dives so seal mounted video cameras were utilized to examine habitat 
use, prey selection, and foraging behavior. Parrish et al. (2000) found that the diurnal pattern of foraging by 
male adults occurred mainly at the 60 m isobath using data collected from Crittercams. A few seals foraged 
at depths of more than 300 m. 

Crittercams also showed that even though much of the seals' time was spent in the shallow areas (<10 m) 
near colonies, the majority of the time searching occurred at greater depths (50 - 60 m). In addition, there 
was evidence of seals consuming prey at deeper depths. Seals targeted habitats that were low-relief and 
areas composed of loose talus fragments provided the best foraging habitat. In this habitat seals are able to 
dislodge the talus fragments and easily locate the prey species. The second most searched habitat type was 
sand-dominated areas where prey was easily accessible. These two habitats were more frequently searched 
and potentially provide higher return of prey for search effort than more complex coral habitats that offer more 
hiding locations for prey species (Parrish et al., 2000). 

Recent studies have focused on characterizing juvenile monk seal habitat use and foraging behavior at French 
Frigate Shoals using Crittercams and TDRs. Juvenile seals forage in the same habitats commonly used by 
adults, but may lack the size and strength to forage as successfully as their adult counterparts (Parrish et al., 
2005). Dive records have indicated that most dives occurred at depths less than 200 m, but occasionally some 
exceeded 200 m. Substantial variability among the pups in depth, duration and temporal patterns of dives was 
noted (NMFS, unpublished data). 

Population Status and Trends 

Current Abundance and Distribution of Populations 

Hawaiian monk seals occur as a single meta-population, with subpopulations distributed among eight NWHI 

locations from Nihoa Island to Kure Atoll, as well as a small and likely growing subpopulation in the MHI. There 

has been variation in the population dynamics between the subpopulations, with differences in environmental 

conditions and levels of human disturbance contributing to this variation (NMFS, 2007; Figure 6.34). 



350 -| 


French Frigate Shoals 






350 -| 




Laysan 


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CD 


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1962 
1965 
1968 
1971 
1974 
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1980 
1983 
1986 
1989 
1992 
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in 


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Figure 6.34. Population trend index (mean beach counts) from individual Hawaiian monk seal subpopulations (~o~ indi- 
cates less reliable historical counts). Source: NOAA PIFSC, unpublished data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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Figure 6.34 (continued). Population trend index (mean beach counts) from individual Hawaiian monk seal subpopulations 
(--o-- indicates less reliable historical counts). Source: NOAA PIFSC, unpublished data. 



Population Trends Across the Northwestern Hawaiian Islands 
The six major subpopulations in the 
NWHI can be divided into the three west- 
ern subpopulations at Kure Atoll, Mid- 
way Atoll and Pearl and Hermes Atoll; 
and the more geographically isolated 
populations found at Laysan and Lisian- 
ski Islands, as well as French Frigate 
Shoals. The higher exchange of indi- 
viduals between the western subpopu- 
lations may contribute to observed simi- 
larities in population dynamics among 
these sites, and these islands may, in 
certain circumstances, be considered a 
single management unit. Table 6.3 illus- 
trates the variation in abundance esti- 
mates between the different islands and 
atolls in 2007. 



Table 6.3. Estimated 2007 monk seal abundance for each population seg- 
ment. Nmin calculated at Mokumanamana and Nihoa Islands according 
to the methods of Wade and Angliss (1997). Source: NMFS, unpublished 
data. 



SITE 


ESTIMATION METHOD 


Nmin 


Nbest 


French Frigate Shoals 


Minimum 


228 


228 


Laysan Island 


Minimum 


209 


209 


Lisianski Island 


Total enumeration 


174 


174 


Pearl and Hermes Atoll 


Minimum 


154 


154 


Midway Island 


Minimum 


65 


65 


Kure Atoll 


Minimum 


105 


105 


Mokumanamana 


Corrected beach counts 


31.4 


42 


Nihoa Island 


Corrected beach counts 


72.2 


78.7 


Main Hawaiian Islands 


Minimum 


88 


88 


Total 




1,126.60 


1,143.70 



The estimated probabilities of sighting an animal is 90% for all years of data at French Frigate Shoals, Laysan 
Island, Midway Atoll and Kure Atoll, approximately 85% at Lisianski Island, and approximately 80% at Pearl 
and Hermes Atoll (Harting, 2002). Therefore, the numbers may underestimate the size of those sub-popula- 
tions by 10%-20%. The methods used for the other population components (Mokumanamana, Nihoa and the 
MHI), while somewhat less accurate, are the best possible under current budget and logistical constraints. 
Because the other population segments represent relatively small proportions of the total population, errors in 
their abundance estimates do not greatly distort the estimated total population size. For example the best esti- 
mate of the total population size in 2005 was 1,247 seals. To determine a minimum population estimate (Nmin) 
for the total population that accounts for the statistical uncertainty in the abundance estimates, as is done for 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



stock assessment reports required by the MMPA (Wade and Angliss, 1997), a combination of enumeration 
and minimum estimates was used. The number of seals identified in 2005 at the main reproductive sites was 
1,065 seals, including 163 pups. Minimum population sizes for Mokumanamana and Nihoa Islands (based on 
the formula provided by Wade and Angliss, 1997) were 34 and 39, respectively, and a total of 77 seals were 
identified in the MHI. Using that procedure, minimum population size estimate for the total population is the 
sum of these estimates, or 1,215 seals. 

Long-term and Recent Population Trends 

Beach counts are used as an index of the population size. Based on the beach counts from the 1950s to 2001 
it appears that the species declined by approximately 50% between the late 1950s and the mid 1970s (Ke- 
nyon, 1973; Johnson et al., 1982). Beach counts of non-pups (juveniles, sub-adults and adults) declined by 
68% between the years 1958 and 2005. 

Based on the very limited window of data availability, the largest counts of monk seals observed at many 
islands were obtained around 1958. Three exceptions to this were French Frigate Shoals, where the maxi- 
mum count was obtained in 1985, Mokumanamana where the maximum was obtained in 1977, and Nihoa 
where the maximum was obtained in 1991. The sum, across all sites in the NWHI, of the maximum counts, 
whenever obtained, totals 1,541, corre- 
sponding, after a very uncertain correc- 
tion, to an estimated population size of 
around 3,000. There is evidence, at a 
minimum, that the available habitat was 
probably capable of supporting at least 
3,000 monk seals in the NWHI plus an 
unknown number in the MHI. 



More recent data indicate that non-pup 
beach counts declined rapidly from 
1985 to 1993, became relatively stable 
for several years, then declined again 
beginning in 1998. Models estimate that 
the total counts declined 8.1% per year 
until 1993 (Carretta et al., 2005). The 
trend in total abundance of Hawaiian 
monk seals is shown in Figure 6.35. A 
log-linear regression of estimated abun- 
dance from 1998 (the first year for which 
a reliable total abundance estimate has 
been obtained) to 2006 estimates that 
abundance declined -3.9% yr - 1 (NMFS, 
unpublished data). 



1500 



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1300 



0) 
O 

c 
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< 

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i i i i i i i i i i 



1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 



Figure 6.35. The long-term combined trend at the main NWHI sites masks 
a diversity of trends within the individual Hawaiian monk seal sub-popula- 
tions. The population dynamics at the different atolls have varied consid- 
erably and current demographic variability among the island populations 
probably reflects a combination of human and environmental/natural influ- 
ences. Source: Gerrodette and Gilmartin, 1990; Polovina et al., 1994; Craig 
and Ragen, 1999; Ragen, 1999. 



The Population Trends by Island and Atoll 

In addition to examining the overall trend in the Hawaiian monk seal population it is also important to examine 
each island and atoll. Each subpopulation exhibits different population dynamics that reflect their unique histo- 
ries and environmental conditions (Figure 6.34). 

Ku re Atoll 

The Hawaiian monk seal population at Kure Atoll has been impacted over the years by human disturbance. 
Beginning with sailors stranded after the ship wreck of the Parker who killed seals for food and continuing with 
disturbance caused by the U.S. Coast Guard (USCG) LORAN station. In the late 1970s NMFS began efforts 
with the USCG to reduce the level of disturbance to the Hawaiian monk seals. USCG decommissioned the 
LORAN station in 1992. Since 1992 state of Hawaii Department of Land and Natural Resources and the Ma- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

rine Mammal Research Program (MMRP) field camps are the only human presence at Kure Atoll during the 
summer months. 

The population dynamics of monk seals have varied with these changes in human disturbance at Kure Atoll 
(Figure 6.34). After the construction of the LORAN station in the late 1950s and early 1960s the seal popula- 
tion declined abruptly. As efforts were made in the 1980s to reduce disturbance to the seals there was a corre- 
sponding increase in pup survival (Gerrodette and Gilmartin, 1990; Gilmartin, pers. comm.). From 1983-2000 
the beach count data shows a 5% increase per year. Since 2000 as seen at other islands and atolls there has 
been high juvenile mortality. The increase in the population until 2000 has been attributed to the reduction of 
human disturbance by USCG regulations and the closure of the LORAN station (Gilmartin et al., 1986). In ad- 
dition 54 immature female seals were released at Kure and reached reproductive maturity by the early 1990s. 
(NMFS, 2007). Recent trends indicate a decline in the Kure population. 

Midway Atoll 

Midway Atoll has also had a history of human disturbance, starting in 1859, which has impacted the Hawaiian 
monk seal population (Figure 6.34). The monk seal population was depleted by the late 1800s and partially 
recovered in the early 1900s. Human activities in the early 1900s included attempts to blast a ship channel, 
installation of a cable station, construction of an airport and other World War II military activities. After World 
War II the human population peaked at 3,500 and was reduced to 250 by 1978 (NMFS, 2007). The 1957-1958 
monk seal surveys recorded a mean of 57 seals but by 1968 only one seal was observed (Kenyon, 1972). 
There were occasional sightings in low numbers during the 1980s (NMFS, 2007). Midway's small monk seal 
population saw increases in the early 1990s as management efforts were increased to reduce disturbance. By 
1988 the U.S. Fish and Wildlife Service (USFWS) was actively participating in the management of wildlife at 
Midway Atoll in conjunction with the U.S. Navy. Since 1988, the USFWS has restricted the numbers of staff and 
visitors to Midway Atoll. Hawaiian monk seal beach counts increased during the 1990s but this was primarily 
due to immigration of individuals from Pearl and Hermes Atoll and Kure Atoll. In addition to immigrants, there 
were increases in total births. In 1996 the Navy transferred Midway Atoll to the USFWS and further measures 
were put in place to reduce disturbance to monk seals. There was a short five year increase in beach counts 
from 1995 - 2000 followed by a decline since 2000. 

Pearl and Hermes Atoll 

Impacts on the seal population at Pearl and Hermes Atoll started in 1859 when a sealing expedition visited the 
atoll. In the late 1920s pearl oysters were harvested and construction occurred on the islands. The U.S. military 
occupied the atoll in 1961 during construction of an observation tower. The atoll is now unoccupied except for 
NMFS summer field camps. 

The Pearl and Hermes Atoll seal population declined by an estimated 90% after the 1950s (Figure 6.34). From 
the mid-1970s to 2000 beach counts have increased. Specifically from 1983 to 2000 counts increased an aver- 
age of 6%. As seen at Kure Atoll and Midway Atoll counts have declined since 2000. 

Lisianski Island 

Lisianski Island is the site of many 1900s ship wrecks and as seen with other wreck sites stranded sailors 
relied on monk seals as well as other species for food. Harvesting expeditions also occurred during this time. 
Today the island is unoccupied except for monk seal field camps. 

Beach counts have declined since the late 1950s and have remained low since then (Figure 6.34). The island 
is estimated to be below the carrying capacity but the cause for low population size is unknown. 

Laysan Island 

The 1857 Hawaiian vessel Manuokawai reported Hawaiian monk seals at Laysan Island. By the 1900s seal 
expeditions and guano miners had nearly extirpated the seal population. For several years the Japanese 
collected eggs and feathers from the island but since this collection was halted in 1915, the island has been 
relatively undisturbed by human presence. The only activities since that time have been survey and scientific 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

expeditions. Beginning in the early 1990s the USFWS has had a small year round field camp on Laysan, and 
NMFS has conducted seasonal field camps on the island. 

As on other islands, Laysan's monk seal population declined in the late 1950s. The population has seen some 
increases but is still below the historic high. Ciguatera is suspected in a mass die off of 50 seals in 1978, but 
was not conclusively proven. In the early 1980s the sex ratio at Laysan was male biased resulting in 45 con- 
firmed deaths due to male aggression from 1982 - 1994. From 1983 - 1994 an average of 4% of the Laysan 
adult females died from injuries related to male aggression (Johanos et al., 1999). Interventions to remove 
subordinate males and correct the sex ratio were successfully undertaken in the mid-90s and have resulted 
in substantially fewer occurrences of male aggression. Even though juvenile survival at Laysan is better than 
French Frigate Shoals there are still concerns about food limitations. There is still not a good understanding of 
the underlying causes or the lack of recovery of the monk seal population at Laysan Island. 

French Frigate Shoals 

French Frigate Shoals did not have the same guano resources that other islands in the NWHI did. As a result, 
the atoll was not mined; however, there is still documentation of harvesting occurring in 1882 of sharks, turtles, 
beche-de-mer and birds. The highest level of use occurred during World War II when a Naval airbase was built 
on Tern Island in 1942. A USCG LORAN station was established and maintained on East Island from 1943 - 
1952. The Naval airbase on Tern Island was decommissioned in 1946 but the USCG maintained the island as 
a LORAN station from 1952-1979. Since the closure of the USCG LORAN station the USFWS has operated a 
field station on Tern Island. 

French Frigate Shoals currently supports the largest monk seal colony in the NWHI (Figure 6.34). The monk 
seal population at East Island and Tern Island were impacted by disturbances caused by the military and 
USCG presence; however, after the closure of the LORAN station in East Island, numbers of monk seals us- 
ing the island increased until the 1980s. After the USCG left Tern Island the presence of seals at the island 
increased until 1989. Since 1989 the French Frigate Shoals population has declined by as much as 75%. 
Juvenile survival rates have declined during this time. Survival in the mid-1980s for weaning to age two was 
as high as 90% but dropped to a low of 8% in 1997. One result of the low survival rates is an imbalance in the 
age structure of the population. The overall population is expected to decline in coming years as fewer females 
reach reproductive age and older females die. 

Mokumanamana and Nihoa Island 

Population monitoring visits to Mokumanamana and Nihoa Islands are infrequent and brief, so enumeration 
is not possible at these sites. Counts of seals at those islands tended to increase from approximately 1970 
to 1990 (Figure 6.34). The increase in counts may have been due to an influx of seals from French Frigate 
Shoals, which was growing at that time. During a seven-day period at Mokumanamana in 1993, 14 tagged 
seals were sighted, all of which had been marked as pups at French Frigate Shoals (Finn and Rice, 1994). 
During the same period, 12 tagged seals were sighted at Nihoa Island, 10 of which were from French Frigate 
Shoals (Ragen and Finn, 1996). 

The Main Hawaiian Islands 

The number of documented monk seal sightings in the MHI increased during the 1990s. Historical abundance 
data for the MHI are limited, as there were no systematic surveys of monk seals conducted prior to 2000. Births 
in the MHI have become more frequent. The known number of annual births in the MHI before and during the 
1990s was usually zero and never exceeded four, but seven births were recorded in 2000 and 12 in 2001 
(Baker and Johanos, 2004). More recently, a minimum of 88 individual seals have been identified in the MHI 
(NMFS, unpublished data). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



MARINE TURTLES 

Covering more than 360,000 km 2 of 
subtropical waters in the North Pacific, 
the Monument provides critical breeding 
and nesting habitat for the green turtle 
(Figure 6.36), and foraging habitat 
and migration pathways for green 
loggerhead, hawksbill and leatherback 
(Figure 6.36) turtles. Although olive ridley 
turtles have not been sighted within the 
Monument, their known distribution in 
the tropical Pacific indicates that they 
likely also use waters of the NWHI 
(Papahanaumokuakea Marine National 
Monument, 2008). These five marine 
turtle species are all protected under 
the ESAdue to threats to beach nesting 
habitat and from incidental bycatch, 
marine debris entanglement, and vessel 
strikes (http://www.nmfs.noaa.gov/pr/ 
species/turtles/). 

Although four marine turtle species have 
been documented in Monument waters, 
the extensive nesting of green turtles in 
the NWHI has allowed for multi-island 
surveys, capture-mark-recapture work 
and long-term population studies of this 
species (Figure 6.37). Based on green 
turtle basking and nesting surveys 
across islands/atolls and over time, the 
species basks throughout the NWHI 
(Figure 6.38), but nests at only a subset 
of the islands (Figure 6.39). 




Figure 6.36. Leatherback turtles (left) and the Hawaiian green turtle (right) 
can be found in the Monument. Photos: NOAA National Marine Sanctuar- 
ies. 




Figure 6.37. Locations of green turtle basking and nesting attempt surveys 
conducted within the NWHI. Map: K. Keller. 



Monitoring Efforts by Island: Data and Methodology 

French Frigate Shoals 

More than 90% of Hawaiian green turtle nesting occurs at French Frigate Shoals, with over 50% of nesting 

occurring on East Island. In addition to intensive study of green turtles at this atoll, annually since 1973, early 

observations were made by military personnel, fishermen and scientists at French Frigate Shoals beginning 

in 1859. Amerson (1971) summarizes all known observations of basking green turtles and nest attempts from 

1859 through 1969. 

The largest data set on green turtles in the NWHI consists of 35 years of research, beginning in 1973, on the 
nesting population on East Island, French Frigate Shoals (Balazs and Chaloupka, 2006). Between 1973 and 
1981, the work was a partnership between the Hawaii Institute of Marine Biology (University of Hawaii) and the 
USFWS, and since 1982 a collaboration between USFWS and the NOAA Pacific Islands Fisheries Science 
Center (PIFSC). From 1973 through the present time, annual surveys have been conducted on the number of 
individual female turtles going ashore each night during the nesting season (Balazs, 1976, 1980; Wetherall et 
al., 1998). Prior to 1996, unique individuals were identified based on double-tagging with external flipper tags; 
since that year, all individuals have been double-tagged with passive integrated transponders. Tagging studies 
have shown that nesting-island site fidelity is very high within the Hawaiian rookery, such that annual nesting 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Legend 

* Basking Sites 
I I Papahanaumol<u3kea Marine National Monument Boundary f 



400 
3 Kilometers 



Figure 6.38. Islands and atolls at which green turtles have been observed 
to bask. Map: K. Keller. 



abundance and trend estimates are not 
confounded by substantial immigration 
or emigration (Dizon and Balazs, 1982; 
Niethammer et al., 1997). 

Annual nesting abundance at East 
Island, French Frigate Shoals, has 
been estimated by the PIFSC using a 
Horvitz-Thompson type estimator: 1ST = 
n/Pj, where NT = number female nest- 
ers in year i, n ; = number of uniquely- 
identified female nesters in year i, and 
p : = probability of sighting a unique fe- 
male that nests in year i. The value p : 
was estimated based upon census data 
from >1,100 nesters during a five-year 
season-long saturation tagging and re- 
sighting program at the atoll from 1988- 
1992. Trends in nester abundance from 
1973-2004 were estimated by Balazs 
and Chaloupka (2004, 2006) using a 
Bayesian smoothing spline regression 
that was fitted to the Horvitz-Thompson 
nest abundance series (Balazs and Ch- 
aloupka, 2004, 2006). 

Other Islands and Atolls 
Data on the number of green turtle pits 
and number of basking turtles at other 
islands and atolls in the northwestern 
chain have been collected in some 
years from 1982 through 2008. Num- 
ber of pits does not indicate a specific 
number of nesting attempts by turtles, 
as each nesting attempt may consist 
of one too many pits; however, pits 
are likely indicative of some level of at- 
tempted nesting on the islands on which 
they have been observed (S. Kubis Har- 
grove, pers. comm.). On Laysan Island, 
NMFS personnel counted the number 
of turtle pits and basking turtles from 
March through June 1982 (Kam, 1986). In 2007, USFWS personnel surveyed the perimeter of Laysan Island 
regularly throughout the green turtle nesting season and monitored active nests for hatching (Payne et al., 
2007). On Lisianski Island, NMFS personnel counted the number of pits dug and number of basking turtles 
through various portions of the nesting season each year from 1982 to 1987, and 2006 and 2007 (Kam, 1985; 
Kam, 1986; Alcorn et al., 1988; Johanos and Withrow, 1988; Westlake and Siepmann, 1988; Kubis, 2008; M. 
Snover, pers. comm.). Similar counts were done at Pearl and Hermes Atoll in 1982, 1990, 1991, 2006, and 
2007 (Kam, 1986; Finn et al., 1993; Kubis, 2008; M. Snover, pers. comm.). At Midway Atoll, the first observa- 
tion of nesting occurred in 2006, and nests were also documented in 2007 and 2008 (John Klavitter, pers. 
comm.). In addition to pit counts at these islands and atolls, in 2006 NMFS personnel quantified the mean 
numbers of basking turtles from May through August at all islands and atolls north of Mokumanamana (B. 
Becker, M. Snover, pers. comm.). 




Figure 6.39. Islands and atolls at which green turtle nesting attempts (pits) 
have been observed. Map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Results and Discussion 
French Frigate Shoals 

Amerson (1971) reports that green turtles were first recorded at French Frigate Shoals by personnel on the 
USS Fenimore Cooper in 1859. In 1882, the crew of the Japanese-owned schooner Ada reported collecting 
1,543 pounds of turtle shell and 47 gallons of turtle oil from approximately 350 turtles. In 1914, green turtles 
were observed basking at French Frigate Shoals by the USS Rainbow's hydrographic survey team, and in 
1923, members of the Tanager Expedition also reported observing turtles and turtle eggs at the atoll. They also 
reported evidence of previous turtle slaughter (Amerson, 1971). 

From the early 1920s through the late 1960s, U.S. Department of Interior, NOAAand the Smithsonian Institu- 
tion personnel reported observing green turtles basking on most of the islands within French Frigate Shoals, 
including Tern, Trig, Whaleskate, Round, East, Gin, Little Gin and Disappearing Islands. The largest num- 
bers of baskers during this time period were reported on East Island in the 1960s, with the highest number 
recorded (86 turtles) in September 1966. Turtle pits were also recorded through the late 1960s on Tern, Trig, 
Whaleskate, Round, East, Gin and Little Gin Islands (Amerson, 1971). 



More recently, NOAA personnel have quantified the number of green turtles basking on all islands within 
French Frigate Shoals during the nesting season. In 2006, an average of 143 turtles was observed basking at 
any one time. 



Results of Balazs and Chaloupka's re- 
search on green turtles nesting at East 
Island, French Frigate Shoals, indicate 
annually variable nesting population siz- 
es, ranging from just under 100 to more 
than 500 female turtles each year from 
1973 to 2004 (Figure 6.40). Estimation 
of nesting population trends indicates 
an increase in annual population size of 
approximately 5.7% per year (95% CI: 
5.3 - 6.1%) over that 32-year time peri- 
od (Figure 6.40; Balazs and Chaloupka, 
2006). 

The increase in green turtle nest- 
ing population size at French Frigate 
Shoals (Figure 6.41) may be attributed 
in part to increased protections under 
the ESA, as harvesting of turtles on land 
or in waters surrounding the Hawaiian 
Islands was prohibited beginning in the 
late-1970s. However, human impacts 
on beach nesting habitat at French Frig- 
ate Shoals also changed dramatically 
over the last half of the 20th century. 
From the late 1930s to early 1950s, the 
U.S. Navy and USCG operated Naval 
Air and Long Range Navigation stations 
on East Island; during that time, turtle 
nesting and basking reportedly greatly 
decreased (Amerson, 1971). After the 
decommissioning of the Naval Air Sta- 
tion and relocation of the USCG LORAN 
station from East Island to Tern Island in 



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Figure 6.40. Trends in green turtle nester abundance at East Island, French 
Frigate Shoals, 1973-2004. Panel A shows time series plot of the Horvitz- 
Thompson estimate of number of female turtles nesting each year over the 
32-year period. Panel B shows estimated long-term trend in nester abun- 
dance derived using Bayesian smoothing spline regression model, which 
was fitted to the Horvitz-Thompson nester series shown in panel A. Red 
curve is mean annual nester abundance. Source: Balazs and Chaloupka, 
2006. 





Figure 6.41. Green turtle laying eggs, French Frigate Shoals, June 2002. 
Photo: NOAA. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

1954, numbers of nest attempts on East 
Island began to increase through the late 
1960s (Amerson, 1971). The number of 
nest at French Frigate Shoals contin- 
ued to increase through 2004 (Balazs 
and Chaloupka, 2006), and during the 
2008 season, an estimated 589 females 
nested at East Island (G. Balazs, pers. 
comm.). 

Other Islands and Atolls 
Numbers of green turtle nesting and 
basking are much lower at the other 
islands and atolls in the northwestern 
chain, relative to French Frigate Shoals. 
The islands of Nihoa and Mokumana- 
mana contain only small areas of bask- 
ing habitat, while Laysan, Lisianski, and the islands of Pearl and Hermes Atoll have small numbers of nesting 
green turtles each year (fewer than 100 turtle pits per island or atoll). Midway and Kure Atolls have relatively 
extensive areas of beach habitat, but only one nest per year has been observed at Midway Atoll since 2006 
(John Klavitter, USFWS, pers. comm.), and nesting has never been observed at Kure Atoll (Kubis, 2008). The 
following sections provide results of pit and basking turtle counts for Nihoa, Mokumanamana, Laysan, Lisian- 
ski, Pearl and Hermes Atoll, Midway Atoll and Kure Atoll for some years from 1982 through 2008. 

Nihoa and Mokumanamana 

The two high islands of the northwestern chain, Nihoa and Mokumanamana, each contain habitat for basking 
and/or nesting, although consisting of only several square meters. There is a small sand beach at Nihoa, but 
only rocky shoreline on Mokumanamana. Green turtles have not been observed nesting on either of these 
islands, but basking has been observed (Kubis, 2008). In 1983, NOAA personnel recorded a mean of 1.5 bask- 
ing turtles per day on Mokumanamana from 24 July through 6 August (Morrow and Buelna, 1985). 

Laysan Island 

In 1982, NOAA personnel on Laysan Island observed 12 green turtle nest excavations, consisting of 45 pits, 
between 25 May and 30 June. In addition, a mean of 2.6 turtles were observed basking each day during the 
period 16 March through 30 June (Kam, 1986). 



In 2005 and 2006, USFWS personnel noted a "few" green turtle pits on Laysan from April through October (C. 
Rehkemper, USFWS, pers. comm.), and in 2006 NMFS personnel recorded 0.2 turtles basking at any one time 
during the nesting season (M. Snover, 
NMFS, pers. comm.). 

In 2007, USFWS located a total of 50 
green turtle nests on Laysan (Figure 
6.42). Of 24 nests that were monitored 
from May through early September, an 
estimated 1,403 eggs were laid, with 
clutch sizes ranging from 55 to 111 
(mean = 87.8, n = 16 nests). Hatching 
success at the 24 monitored nests was 
85.6%. Incubation periods ranged from 
60-75 days (mean = 67.5), with incuba- 
tion period shortening over the course 

of the season (Payne et al., 2007). 

Figure 6.42. Green turtle nest on Laysan Island, 26 June 2007. Photo: US- 
FWS. 





A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Lisianski Island 

NMFS personnel counted turtle pits and basking green turtles on Lisianski Island during parts of the nesting 
season in some years between 1982 and 2007. Results of these counts indicate a range of 15 to 47 pits ob- 
served per year, and a mean of 2.0 to 5.4 turtles basking on the island per day (Table 6.4). Figure 6.43 indi- 
cates the locations of pits observed on Lisianski Island in 1982 and 1983. 

Table 6.4. Observations of pits and basking green turtles on Lisianski Island, 1982-2007. 



YEAR 


DATES 


OBSERVATIONS 


REFERENCE 


1982 


May -Aug 


23 excavation sites consisting of 47 pits 


Kam (1986) 


1982 


8 Jul -13 Sep 


Mean of 5.4 baskers/day 


Kam (1986) 


1983 


31 May - 9 Aug 


19 pits (Figure 6.43) 


Kam (1985) 


1984 


2 Jul -6 Aug 


91 pits 


Alcorn et al. (1988) 


1985 


17 Jun-20 Jul 


78 pits 


Alcorn et al. (1988) 


1986 


5 -26 Aug 


15 pits 


Westlake and Siepmann (1988) 


1987 


1-4 Jun; 5 -29 Aug 


34 pits 


Johanos and Withrow (1988) 


1987 


7 -27 Aug 


Mean of 2.0 baskers/day 


Johanos and Withrow (1988) 


2006 


May -Aug 


Mean of 2.9 baskers/day 


Melissa Snover, NMFS (pers. comm.) 


2007 


May- Jun 


Nests observed but not counted 


Kubis (2008) 




Figure 6.43. Locations of green turtle pits on Lisianski Island May -August 
1982 and May - August 1983. Sources: Kam, 1986 and!985. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Pearl and Hermes Atoll 
NMFS personnel observed turtle pits 
and basking green turtles at Pearl and 
Hermes Atoll (on North, Southeast and 
Seal Kittery Islands) during parts of the 
nesting season in some years between 
1982 and 2007 (Kam, 1986; Finn et al., 
1993; Kubis, 2008). Count data indicate 
a range of one to 13 pits observed per 
year, and a mean of 10.0 to 12.6 turtles 
basking on North and Southeast Islands 
per day (Table 6.5). Figure 6.44 indi- 
cates the locations of pits observed on 
North and Southeast Islands in 1982. 

Midway Atoll 

One green turtle nest per year has been 
found at Midway Atoll since 2006. In 
2006, a single nest was located on Spit 
Island, and in 2007 and 2008 one nest 
per year was found on Sand Island (J. 
Klavitter, USFWS, pers. comm.). All 
three nests (2006, 2007, and 2008) 
hatched successfully. The nest laid in 
2008 was estimated to have contained 
89 eggs, 65 of which hatched success- 
fully (J. Klavitter, USFWS, pers. comm.; 
Figures 6.45 and Figure 6.46). In 2006, 
NMFS personnel observed 4.5 basking 
turtles at any one time at Midway Atoll 
(M. Snover, pers. comm.). 

Kure Atoll 

Nesting by green turtles has never been 
observed at Kure Atoll (Kubis, 2008). 
Biological monitoring and research on 
other species has been conducted by 
NMFS and Hawaii Department of Land 
and Natural Resources personnel at 
Kure over a period of many years, so 
it is unlikely that nesting has occurred 
but has remained unobserved. Green 
turtles have been observed basking at 
Kure Atoll in small numbers. In 2006, 
NMFS personnel observed 0.7 bask- 
ing turtles at any one time during a total 
of eight observation days (M. Snover, 
NMFS, pers. comm.). 



Table 6.5. Observations of pits and basking green turtles at Pearl and 
Hermes Atoll, 1982-2007. 



YEAR 


DATES 


ISLAND 


OBSERVATIONS 


REFERENCE 


1982 


5 Jul 


North Island 


13 pits 
(Figure 6.44) 


Kam (1986) 


1982 


2-5 Jul 


Southeast 
Island 


9 pits 
(Figure 6.44) 


Kam (1986) 


1982 


2-6 Jul 


North and 
Southeast 


Mean of 10 
baskers/day 


Kam (1986) 


1990 


8-9 Jun 


North Island 


2 pits 


Finn et al. (1993) 


1991 


1 Aug - 13 
Sep 


Southeast 
Island 


lpit 


Finn et al. (1993) 


2006 


May - Aug 


Not specified 


12.6 baskers/day 


Melissa Snover, 
NMFS (pers. comm.) 


2007 


May- Jun 


Not specified 


Nests observed 
but not counted 


Kubis (2008) 



B 




Figure 6.44. Panel A: locations of green turtle pits observed at Pearl and 
Hermes atoll in 1982. Panel B indicates locations of pits on Southeast Is- 
land; panel C indicates pit locations on North Island. Source: Kam, 1986. 




Figure 6.45. Green turtle tracks and nest on Sand Island, Midway Atoll, May 
2008. The left panel shows turtle tracks and the right panel shows an active 
nest. Photos: T Summers, USFWS. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Conclusions and Management 
Action 

Green turtles have been observed 
basking at all of the islands and atolls 
within the Monument, but nesting at- 
tempts have been limited to the islands 
and atolls from French Frigate Shoals to 
Midway Atoll. As previously described, 
more than 90% of green turtle nesting 
occurs at French Frigate Shoals, with 
a range of approximately 100 to 550 
females digging approximately 450 to 
2,500 nests each year (Table 6.6). In 
contrast, between 15 and 91 pits have 
been observed on Laysan and Lisian- 
ski Islands in any one year, 1-13 pits 
per island at Pearl and Hermes Atoll 
per year, and only one nest per year at 
Midway Atoll between 2006 and 2008 
(Table 6.5). As the number of nesting 
females on East Island, French Frig- 
ate Shoals has increased over the last 
three decades, the numbers of nests 
at Laysan, Lisianski, and Pearl and 




Figure 6.46. Location of green turtle nest on Sand Island, Midway Atoll, in 
2008. Source: J. Klavitter; map: K. Keller. 



Hermes Atoll may have also increased over time. Additional periodic counts of pits and/or mark-resighting of 
nesting females at those locations in future years will help to more precisely estimate the subpopulation sizes 
of nesting green turtles throughout the Monument. 

Table 6.6. Numbers of green turtle nest pits observed on all islands and atolls in the NWHI between 1982 and 2008. 
Dash (-) indicates data not collected or not available; plus sign (+) indicates observed pits but not counted; and zero (0) 
indicates true count of zero. 



ISLAND/ATOLL 


1982 


1983 


1984 


1985 


1986 


1987 


1990 


1991 


2006 


2007 


2008 


Nihoa 



































Mokumanamana 



































French Frigate Shoals: East 
Islandt 


585* 


158* 


896* 


729* 


311* 


644* 


675* 


482* 


1,904* 


1,566* 


2,430* 


Laysan 


45 


- 


- 


- 


- 


- 


- 


- 


- 


50 


- 


Lisianski 


47 


19 


91 


78 


15 


34 


- 


- 


- 


+ 


- 


Pearl and Hermes Atoll: 
North Island 


13 


- 


- 


- 


- 


- 


2 


- 


- 


+ 


- 


Pearl and Hermes Atoll: 
Southeast Island 


9 


- 


- 


- 


- 


- 


- 


1 


- 


+ 


- 


Midway Atoll: 


- 


- 


- 


- 


- 


- 


- 


- 


- 


- 


- 


Spit Island 


























1 








Midway Atoll: 
























Sand Island 





























1 


1 


Kure Atoll 



































tPits have also been observed but not counted in all years on Tern, Trig, Gin and Little Gin Islands at French Frigate Shoals. 

*Numbers of pits at East Island, French Frigate Shoals, are estimates based upon numbers of nesting females (Stacy Kubis Har- 
grove, pers. comm.). Individual turtles lay three to six (mean = 4.5) clutches per season (http://www.fpir.noaa.gov/PRD/prd_green_ 
sea_turtle.html), so estimates of number of nest pits was obtained by multiplying the number of nesting females per year by 4.5. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Papahanaumokuakea Marine National Monument's Draft Management Plan (PMNM, 2008) describes a strat- 
egy and three activities to manage green turtle habitat within the Monument, based upon the Recovery Plan 
for U.S. Pacific Populations of the Green Turtle (Chelonia mydas; NMFS and USFWS,1998). Strategy TES-3 
of the Monument's Draft Management Plan is to "Ensure that nesting populations of green turtles at source 
beaches are stable or increasing over the life of the plan" (the life of the plan being 15 years). The first activity 
planned to achieve this strategy includes the continuation of data collection to monitor nesting turtles on East 
Island, French Frigate Shoals (with the largest numbers of nesting turtles in the Monument), and the periodic 
reassessment of the distribution of nesting activity on the other islands and atolls within the NWHI (PMNM, 
2008). The second activity to achieve Strategy TES-3 is the protection and management of nesting habitat, in- 
cluding prevention of introduction of mammalian predators such as rats, reduction of artificial light near nesting 
beaches, prohibition of habitat alteration, and the regulation of human access and activities. Management ac- 
tions to delay habitat loss due to sea level rise are also advised, but specific activities related to the slowing of 
climate-change-induced habitat loss are not described (PMNM, 2008). Finally, Strategy TES-3 will be attained 
by protecting and managing foraging areas and migration routes within the Monument, including identification 
and mapping of these areas, and management of vessel transit and discharge, and minimization of the intro- 
duction of contaminants (PMNM, 2008). 



EXISTING DATA GAPS 

It is important to develop and regularly update a database of population structure and dynamics for protected 
species. The database will help managers make effective decisions and determine the effects of previous deci- 
sions and events (e.g., climate events, management decisions, research programs, disease outbreaks, etc.). 
Specific opportunities include research to improve the understanding of: 

The essential habitats and ecological requirements of protected species, to minimize anthropogenic 
threats and the effect of catastrophic events; 

The diet and foraging behavior of the Hawaiian monk seals throughout different life stages in order to 
understand the effect of food availability on the population; 

Time budgets, diving, and movement characteristics and energetics of the Hawaiian monk seal, stratified 
by representative sub-populations, age, and sex classes; 

An appropriate and sensitive assay for biotoxins and metabolites in tissue of monk seals and prey spe- 
cies; 

The effects of climate change on nesting sites of protected species, e.g., the effect of sea level rise on 
nesting sites of the green sea turtle and Hawaiian monk seal; 

The Allee effect (i.e., that for smaller populations, the reproduction rates and survival of individuals de- 
crease) and thresholds for phase shift; 

Cetacean presence and behavior in the NWHI at different times of the year; and 

The presence/absence of other turtle species and nesting sites at other locations in the NWHI. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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NOAA. http://www.fpir.noaa.gov/PRD/prd_green_sea_turtle.html 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Seabirds 

Kaylene E. Keller 1 , Angela D.Anders 2 , Scott A. Shaffer 3 , Michelle A. Kappes 3 , Beth Flint 4 and Alan Friedlander 56 



INTRODUCTION 

The Northwestern Hawaiian Islands 
(NWHI) provide habitat for an estimated 
5.5 million nesting seabirds represent- 
ing 22 different species (see Figures 
7.1 and 7.2 and Fefer et al., 1984). For 
many of these species, the Monument 
provides the majority of available nest- 
ing habitat. Nesting occurs throughout 
the year, varying by species and within 
species, and by annual food availability 
(Harrison, 1990). The largest breeding 
populations occur at Midway Atoll, Lay- 
san Island and Nihoa (USFWS, 2005). 
While the Monument protects seabird 
breeding and nesting habitat, the for- 
aging ranges for most of these species 
extend beyond the Monument's bound- 
aries, with foraging distances ranging 
from 3 km to several thousand kilome- 
ters (e.g., Fernandez et al., 2001 ). Some 
seabird species occur year-round in the 
Monument, while others migrate to oth- 
er parts of the Pacific when not breed- 
ing. Juvenile birds may also remain at 
sea for several years before returning 
to their breeding colonies inside Monu- 
ment water. Overall, the NWHI provide 
high-quality breeding habitat, with low 
predation risk and low disturbance con- 
ditions (Table 7.1). 




Figure 7.1. A Red-footed Booby. Red-footed Boobies nest on all islands 
and atolls in the NWHI. Photo: J. Watt. 







Figure 7.2. Brown Noddy Terns and Brown Boobies at Pearl and Hermes 
Atoll. Photo J. Watt. 



The distribution of seabirds within the 
Monument reflects, to some extent, the 
nesting habitat currently available on 

the islands (Tables 7.2 and 7.3). Nesting distribution has also been affected by human disturbance in the late 
19th and 20th centuries, including egg and feather hunting, and destruction of habitat due to military activities 
during and after World War II. For example, populations of many of the seabird species on Laysan Island are 
likely still recovering after the devegetation of the island by guano miners and hunter-introduced feral rabbits in 
the early 1 900s (Ely and Clapp, 1 973). Further habitat losses have occurred from the introduction of non-native 
plants like golden crown-beard (Verbisena encelioides) and rodent pests like rats and mice. Numbers of nest- 
ing adults of some species are also still increasing on Tern and East Islands at French Frigate Shoals after the 
decommissioning of Naval and Coast Guard stations following World War II (Amerson, 1971). 



1 . NOAA/NOS/ONMS/Papahanaumokuakea Marine National Monument 

2. Clancy Environmental Consultants, Inc. 

3. University of California Santa Cruz 

4. U.S. Fish and Wildlife Service 

5. NOAA/NOS/NCCOS/CCMA Biogeography Branch 

6. The Oceanic Institute 




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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 
Table 7.3. Distribution of seabirds in the Pacific, Main Hawaiian Islands (MHI) and the NWHI. Source: USFWS, 2005. 



COMMON NAME 


SCIENTIFIC NAME 








Procellariiformes (Albatrosses, Petrels and Shearwaters) 


Black-footed Albatross 


Phoebastria nigripes 


B 


B 


B 


Laysan's Albatross 


Phoebastria immutabilis 


B 


B 


B 


Short-tailed albatross 


Phoebastria albatrus 


b 






Hawaiian Petre 


Pterodroma sandwichensis 




B 




Herald Petrel 


Pterodroma arminjoniana 






B 


Tahiti Petrel 


Pterodroma rostrata 






B 


Bonin Petrel 


Pterodroma hypoleuca 


B 


Ex 


B 


Phoenix Petrel 


Pterodroma alba 






Ex 


Bulwer's Petrel 


Bulweria bulwerii 


B 


B 


B 


Wedge-tailed Shearwater 


Puffinus pacificus 


B 


B 


B 


Christmas Shearwater 


Puffinus nativitatis 


B 


B 


B 


Newell's Shearwater 


Puffinus auricularis newelli 




B 1 




Audobon's Shearwater 


Puffinus Iherminieri 






B 


Band-rumped Storm-Petrel 


Oceanodroma castro 




B 


B 


Black Storm-Petrel 


Oceanodroma melania 




B 




Tristram's Storm-Petrel 


Oceanodroma tristrami 


B 




B 


Polynesian Storm-Petrel 


Nesofregetta fuliginosa 






B 


Pelecaniformes (Tropicbirds, Boobies and Frigatebirds) 


White-tailed Tropicbird 


Phaethon lepturus dorothea 


B 


B 


B 


Red-tailed Tropicbird 


Phaethon rubricauda melanorhynchos 


B 


B 


B 


Masked Booby 


Sula dactylatra personata 


B 


B? 


B 


Brown Booby 


Sula leucogaster plotus 


B 


B 


B 


Red-footed Booby 


Sula sula rubripes 


B 


B 


B 


Great Frigatebird 


Fregata minor palmerstoni 


B 


b 


B 


Lesser Frigatebird 


Fregata ariel ariel 






B 


Charadriiformes (Terns and Noddies) 


Little Tern 


Sterna albifrons sinensis 


B 




B 


Gray-backed Tern 


Sterna lunata 


B 


B? 


B 


Bridled Tern 


Sterna anaethetus 






B? 


Sooty Tern 


Sterna fuscata oahuensis 


B 


B? 


B 


Brown Noddy 


Anous stolidus pileatus 


B 


B 


B 


Black Noddy 


Anous minutus marcusi 


B 


B 


B 


Blue Noddy 


Procelsterna cerulea saxatilis 


B 


B? 


B 


White Tern 


Gygis alba alba 


B 


B 


B 


Note: 1 Endemic 

Abbreviations: B = breeding; b = unsuccessful breeding attempts; B? = breed' 


ng suspected; Ex 


= extirpated bree 


ders 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




PROCELLARIIFORMES (ALBATROSSES, PETRELS AND SHEARWATERS) 

Laysan Albatross {Phoebastria immutabilis) 
Laysan Albatrosses are a relatively small 
albatross species, with a body length of 
79-81 cm, a wingspan of 195-203 cm, 
and a mean body mass of 2.4 kg (Whit- 
tow, 1993; Suryan et al. 2008; Figure 
7.3). Like Black-footed Albatrosses (P. 
nigripes) and other Procellariiformes, 
males are slightly larger than females 
but feather plumage is identical for both 
sexes. Laysan Albatrosses have a white 
head, neck, and underparts, and sooty 
brown upper wings and trailing edges of 
under wings, and back. The legs, feet, 
and bill are pink but the bill also has a 

greenish tip (Whittow, 1993). Rgure ?3 Laysan A i bat ross, Tern Island and French Frigate Shoals. Pho- 

to: C. Gregory. 
When breeding, Laysan Albatrosses 
nest on all of the islands and atolls of 
the NWHI chain, and at discrete colo- 
nies on Kauai and Oahu, Torishima Is- 
land, Japan (Kurata, 1978), and off of 
the west coast of Baja, Mexico (USFWS, 
2005). A pair was also documented to 
have successfully bred on Wake Island 
in 2001 (USFWS, 2005; Figure 7.4). In 
the NWHI, egg laying and incubation 
are generally synchronous, occurring 
from November to January. Chicks are 
reared from late January to mid-July 
and fledge in mid to late July (USFWS, 
2005). Most birds breed every year but 
will occasionally skip a breeding event 
(Fisher, 1976). 

In the NWHI, predator control, including 
rat eradication, has reduced some of the 
threats to Laysan Albatrosses, but ex- 
isting impacts to nesting habitat include 
the invasive wildflower golden crown- 
beard (Verbesina encelioides), other 
non-native plant species, obstacles to 

flying (USFWS, 2005), and lead toxicity in chicks at Midway from ingestion of lead paint chips from dilapidated 
buildings (Finkelstein et al., 2003). Threats at sea include the ingestion of marine debris (e.g., plastics), sea 
level rise and long-line bycatch (USFWS, 2005). 




Figure 7.4. Laysan Albatross nesting sites and foraging areas in the NWHI. 
Source: USFWS, unpub. data; map: K. Keller. 



When at sea, tracking studies reveal that Laysan Albatrosses are found throughout the North Pacific (including 
the Bering Sea) but generally range between 30°N and 55°N (Harrison, 1987; Fernandez et al., 2001 ; Hyren- 
bach et al., 2002; Shaffer et al., 2005). During the incubation period, adults conduct foraging excursions lasting 
10-30 days and may travel over 2,000 km from the nesting colony (Kappes et al., in review, Figures 7.4 and 
7.5). Albatrosses studied across several consecutive years (2002-2006) show that habitat use changes across 
years (Figure 7.5) but the environmental cues that putatively influence foraging effort remain the same across 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

years studied. That is, sea surface temperature (SST) and primary productivity levels appear to be the most 
important predictors of searching/foraging activity (Kappes et al., in review). During the chick-brooding period, 
adults are constrained by the need to provision their chick frequently so foraging excursions last only one to 
three days on average and range from the colony is typically less than 400 km (Kappes et al., unpublished 
data). When chicks are large enough to defend themselves and are thermally independent, both parents are 
able to forage simultaneously. During this period (i.e., chick-rearing), adults conduct both long and short dura- 
tion foraging trips with some trips extending northward to the Aleutian Islands (Fernandez et al., 2001 ; Hyren- 
bach et al., 2002). When breeding is complete, adult Laysan Albatrosses depart the breeding colonies for cool 
waters of the Central and Western North Pacific including the Bering Sea and Aleutian Islands (Shaffer et al., 
submitted; Figure 7.5). Here, birds remain for most of the summer months while undergoing molt and recover- 
ing from breeding. Albatrosses return to the colonies in mid- to late-November to breed again. It is important 
to note the this species has only been tracked from a few of their breeding colonies, thus at sea distribution is 
not fully characterized. Laysan Albatrosses are surface feeders, with a diet consisting of squid, crustaceans, 
fish and flying fish eggs. At least 50% of the diet is composed of squid (USFWS, 2005). 



Laysan albatross 

Phoeha&ria nnmiiiabilis 



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Figure 7.5. Laysan Albatross utilization distribution during brooding, Incubation and postbreedlng. Source: Shaffer et al., 
In review; maps: R. Clark. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Black-footed Albatross {Phoebastria nigripes) 
Black-footed Albatrosses are slightly 
larger than Laysan Albatrosses, with a 
body length of 64-74 cm, a wingspan of 
193-216 cm, and a mean body mass of 
2.8 kg (Awkerman et al., 2008; Suryan 
et al., 2008; Figure 7.6). Like Laysan 
Albatrosses, males are slightly larger 
than females but plumage is identical 
between the sexes. Black-footed Al- 
batrosses are dusky-brown with white 
fringes around the base of the bill, un- 
der the eye, under the tail, and over the 
base of the tail (Harrison, 1 987). 



Ninety eight percent of the breeding 
population occurs within the Monu- 
ment. The species nests on all of the 
islands and atolls in the NWHI, with the 
majority of pairs nesting at Midway Atoll 
and on Laysan Island (USFWS, 2005; 
Figure 7.7). A few thousand (Awkerman 
et al., 2008) Black-footed Albatrosses 
also breed on discrete colonies in Jap- 
anese Islands, and several pairs have 
been observed prospecting at Guadal- 
upe Island, Baja, Mexico. Within breed- 
ing colonies, Black-footed Albatrosses 
nest synchronously, with all females 
laying eggs within a few-week period. 
Throughout the Hawaiian archipelago, 
egg-laying and incubation generally oc- 
curs from October through December, 
and chicks are reared from January to 
June, with fledging occurring in mid- 
June (Awkerman et al., 2008). Historical 
breeding areas include the Hawaiian ar- 
chipelago and Marshall, Johnston, and 
Torishima Islands (Harrison, 1987). 




Figure 7.6. Black-footed Albatross, Tern Island and French Frigate Shoals. 
Photo: C. Gregory. 




Legend 

■ Black-footed Albatross Nesting Sites 

: Black-fooled Albatross Estimated Foraging Range (700Km) N 

I Papahanaumokuakea p\ 

Marine National Monument 
500 1,000 



Figure 7. 7. Black-footed Albatross nesting sites and foraging areas in the 
NWHI. Source: USFWS, unpub. data; map: K. Keller. 



Within the Monument, many threats 

have been reduced over the years, but 

existing threats include ingestion of plastics, long-line bycatch, and sea level rise (USFWS, 2005; Lewison and 

Crowder, 2003). The Monument provides protection for breeding and nesting habitat, but only a small portion 

of the foraging range remains protected because of the wide ranging foraging behavior. 



When at sea, tracking studies reveal that Black-footed Albatrosses are found throughout the North Pacific 
(partially the Bering Sea) but generally they range between 25°N and 50°N (Harrison, 1987; Fernandez et 
al., 2001 ; Hyrenbach et al., 2002; Shaffer et al., 2005). During the incubation period, adults conduct foraging 
excursions lasting 10-20 days in duration and may travel over 1,500 km from the nesting colony (Kappes et 
al., in review; Figures 7.7 and 7.8). Adults tend to head north-northeast of the breeding islands to warmer wa- 
ters along the southern edge of the North Pacific Current. Like Laysan Albatrosses, Black-footed Albatrosses 
studied across multiple consecutive years (2002-2006) use slightly different habitat across years (Figure 7.8) 
but the environmental cues that putatively influence foraging effort remain the same across years studied. That 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

is, SST and primary productivity levels appear to the most important predictors of searching/foraging activity 
(Kappes et al., in review). During the chick-brooding period, adult Black-footed and Laysan Albatrosses expe- 
rience the greatest overlap in their distribution because adults are constrained by the need to provision their 
chick frequently; thus foraging excursions last only one to three days on average and ranges are typically less 
than 300-400 km from the colony (Kappes et al., unpublished data). During the chick-rearing period, adults 
conduct both long and short duration foraging trips with some trips extending to the west coast of the U.S. and 
Canada (Fernandez et al., 2001 ; Hyrenbach et al., 2002). After breeding, adult Black-footed Albatrosses leave 
the breeding colonies for rich productive waters of the California Current and Aleutian Islands (Hyrenbach 
et al., 2006; Fischer, 2008; Shaffer et al., submitted; Figure 7.8). Thus, the spatial overlap with Laysan Alba- 
trosses during the non-breeding period can be relatively minor (<10%; Shaffer et al., submitted). Black-footed 
Albatrosses remain along the West Coast of the U.S. or Aleutian Islands for most of the summer months while 
undergoing molt and recovering from breeding. Albatrosses return to breeding colonies in mid- to late-October 
to breed again. It is important to note the this species has only been tracked from a few of their breeding colo- 
nies, thus at sea distribution is not fully characterized. The species is a surface feeder, with a diet including 
fish eggs, squid, crustaceans, fish, and zooplankton. Forty percent of the diet is composed of flying fish eggs 
(USFWS, 2005). 

















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Figure 7.8. Black-footed Albatross utilization distribution during brooding, incubation and postbreeding. Source: Shaffer 
et al., in review; maps: R. Clark. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Short-tailed Albatross {Phoebastria albatrus) 
Short-tailed Albatrosses are larger than 
Laysan and Black-footed Albatrosses, 
with a mean wingspan of 237 cm and 
mean body mass of 4.3 kg (Brooke, 
2004; Suryan et al., 2008). Short-tailed 
Albatrosses are similar in appearance 
to Laysan Albatrosses, but are larger, 
with a heavier bright pink bill, and with 
a yellow wash on the white plumage of 
the head and neck (Harrison, 1990; Fig- 
ure 7.9). 



Short-tailed Albatrosses historically 
ranged throughout the North Pacific, but 
current breeding sites include only Tor- 
ishima and Minami-kojima, Japan (USF- 
WS, 2005). Short-tailed Albatrosses pe- 
riodically do attempt to breed at Midway 
Atoll, but there are no documented ac- 
counts of successful nesting (USFWS, 
2005). Eggs are laid from October to 
November and fledging occurs in June. 




Figure 7.9. Short-tailed Albatross. Photo: J. Lloyd. 



The Short-tailed Albatross is listed as Endangered under the Endangered Species Act (ESA) throughout its 
range. Within the Monument, threats are minimal relative to other potential colony sites, and management ac- 
tions are taking place to encourage nesting at Midway Atoll. Tracking studies show that these albatrosses gen- 
erally remain in the Western Pacific when breeding, and then move further north into the Bering Sea and along 
the Aleutian Islands (Suryan et al., 2006). Based on the large body size of Short-tailed Albatrosses compared 
to Laysan and Black-footed albatrosses, Short-tailed Albatrosses would appear to have a more restricted 
range that occurs within regions of stronger winds and larger wave heights (Suryan et al., 2008). 

Bonin Petrel {Pterodroma hypoleuca) 
Bonin Petrels are 30 cm long with a 
wingspan of 63-71 cm and a mean body 
mass of 204 g. Plumage on the upper 
parts is blue- to silver-gray, with a sooty 
head and neck, and white forehead, 
chin and throat. Upper wings are gray 
with black primaries, and the underwing 
is white with black margins (Seto and 
O'Daniel, 1999; Figure 7.10). 

Bonin Petrels are found throughout the 
western north Pacific and breed in the 
NWHI and on Volcano and Bonin Islands 
in Japan. There are no breeding colo- 
nies of Bonin Petrels in the Main Hawai- 
ian Islands (MHI; Harrison, 1987). 

In the NWHI, Bonin Petrels nest at 

French Frigate Shoals, Laysan Island, 

Lisianski Island, Pearl and Hermes Atoll, Midway Atoll and Kure Atoll. Gardner Pinnacles may also provide 

nesting habitat. Lisianski Island, Laysan Island and Midway Atoll support the largest colonies in the archipela- 




Figure 7. 1 0. Bonin Petrel. Photo: C. Gregory. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



go (Figure 7.11). In the NWHI, eggs are 
laid from December to March and chicks 
are reared from February to May. 

Bonin Petrels are nocturnal surface 
feeders, with a diet consisting mainly of 
fish (USFWS, 2005). 

Habitat loss from erosion and invasive 
plant species are some of the main 
threats to the populations in the NWHI. 
Historically, introduced Polynesian rats 
posed a serious threat to Bonin Petrel 
populations, but rats were extirpated in 
the NWHI by the year 2000. The U.S. 
Fish and Wildlife Service (USFWS) has 
also modified light sources at Midway 
Atoll and French Frigate Shoals to re- 
duce impacts to nocturnal seabird spe- 
cies, including the Bonin Petrel. 



Bulwer's Petrel (Bulweria bulwerii) 
The Bulwer's Petrel is 26 cm long with 
a 67 cm wingspan (Harrision, 1987). 
Plumage is sooty brown, with a light- 
er brown face and chin (Megysi and 
O'Daniel, 1997; Figure 7.12). 

The Bulwer's Petrel is a wide-ranging 
species occurring in the tropical and 
subtropical waters of the Pacific, Atlan- 
tic and Indian Oceans (Harrision, 1987). 
In the Pacific, Bulwer's Petrels breed in 
the Phoenix, Marquesas, Bonin, Volca- 
no and Hawaiian Islands, including the 
NWHI and MHI (USFWS, 2005). From 
the little that is known about the spe- 
cies' at-sea distribution, it appears that 
the Hawaiian populations travel to the 
central and eastern Pacific during the 
non-breeding period (USFWS, 2005). 




Figure 7.11. Bonin Petrel nesting sites in the NWHI. Source: USFWS, un- 
pub. data; map: K. Keller. 




Figure 7. 12. Bulwer's Petrel. Photo: USFWS. 



In the NWHI, Bulwer's Petrels have 

been found to breed at Nihoa Island, Mokumanamana, French Frigate Shoals, Laysan Island, Lisianski Island, 
Pearl and Herrmes Atoll and Midway Atoll (Figure 7.13). The largest breeding colony occurs on Nihoa Island. 
Bulwer's Petrels lay eggs in the NWHI between May and July. Chicks are present from July to October, and 
fledging occurs by early October. 

Bulwer's Petrels are nocturnal surface feeders, with a diet composed of fish, squid, crustaceans and sea- 
striders. Most of the prey species are bioluminescent and are found in upwelling areas (USFWS, 2005). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



As seen with many of the seabirds in the 
NWHI, Bulwer's Petrels suffered losses 
to Polynesian rat and ant introductions. 



Wedge-tailed Shearwater 
(Puffin us pacificus) 
The Wedge-tailed Shearwater is 43 cm 
long, with a wingspan of 101 cm (Har- 
rison, 1987; Figure 7.14), and a mean 
body mass of 390 g (Whittow, 1997). 
Light-morph shearwaters are grayish 
brown above, with white underparts, 
while dark-morph individuals are sooty 
brown above and below (Whittow, 
1997). 

Wedge-tailed Shearwaters occur 
throughout the tropic and subtropical 
Pacific and Indian Oceans, including the 
NWHI (Harrison, 1987). The Hawaiian 
populations most likely migrate to the 
Equatorial Countercurrent and east dur- 
ing the non-breeding period (USFWS, 
2005). This species aggregates into 
large multi-species feeding flocks, often 
associated with subsurface predators 
(e.g., tuna and dolphin). 

Wedge-tailed Shearwaters nest on all 
of the islands and atolls in the NWHI 
(Figure 7.15). The largest colonies with- 
in the archipelago are at Nihoa Island, 
Laysan Island and Lisianski Island. In 
the NWHI, eggs are laid from June to 
August and chicks are present from Au- 
gust to December, with the majority of 
fledging occurring in November. 

Wedge-tailed Shearwaters use contact 
dipping for feeding, and their diet in the 
Hawaiian Archipelago consists mainly 
of larval goatfish, flying fish, squirrelfish 
and squid (USFWS, 2005). 




Figure 7. 13. Bulwer's Petrel nesting sites in the NWHI. Source: USFWS, 
unpub. data; map: K. Keller. 




Figure 7. 14. Wedge-tailed Shearwater light and dark morphs, Tern Island, 
French Frigate Shoals. Photo: USFWS. 




Figure 7. 15. Wedge-tailed Shearwater nesting sites and foraging areas in 
the NWHI. Source: USFWS, unpub. data; map: K. Keller. . 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Christmas Shearwater {Puffinus nativitatis) 
Christmas Shearwaters are 36 cm long 
with a 76 cm wingspan (Harrison, 1987), 
and a mean body mass of 354 g (Seto, 
2001). Plumage is dark brown through- 
out, with underparts slightly paler than 
upperparts (Seto, 2001 ; Figure 7.16). 

Christmas Shearwaters occur in the 
tropical and subtropical Pacific (Harri- 
son, 1987). The species nests on all of 
the islands and atolls in the NWHI, with 
the largest breeding colonies occurring 
on Laysan Island and Lisianski Island 
(USFWS, 2005; Figure 7.17). Females 
lay eggs from April through July, and 
chicks are reared from June to October. 



Christmas Shearwaters use pursuit and 
plunge feeding behavior, with a diet 
composed of fish (goatfish, flying fish 
and scad) and squid (USFWS, 2005). 

Nesting habitat degradation due to the 
spread of non-native invasive plant spe- 
cies is the most significant threat to the 
species in the NWHI. 




Figure 7.16. Christmas Shearwaters, 
Shoals. Photo: D. Wright. 



Tern Island and French Frigate 




Legend 



Christmas Shearwater Nesting Sites 



| | Papahanaumokuakea 

Marine National Monument 

20'] 400 55! 



Figure 7.17. Christmas Shearwaters nesting sites in the NWHI. Source: 
USFWS, unpub. data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Tristram's Storm-petrel {Oceanodroma tristrami) 
The Tristram's Storm-petrel is the small- 
est Procellariiformes species in the 
NWHI. It averages 24 cm in length, with 
a 56 cm wingspan and a mean body 
mass of 77-97 g. The species is brown- 
ish gray throughout, with a buffy brown 
wing bar across the upper wing (Slot- 
terback, 2002; Figure 7.18). 

The Tristram's Storm-petrel range in- 
cludes the subtropical central and west- 
ern Pacific and waters off of Japan. 
Breeding colonies occur in the NWHI, 
but there is no evidence of colonies in 
the MHI (USFWS, 2005). 

The species is found on Nihoa Is- 
land, Mokumanamana, French Frigate 
Shoals, Laysan Island, Lisianski Island, 
and Pearl and Hermes Atoll. The larg- 
est colonies are at Nihoa Island, Laysan 
Island, and Pearl and Hermes Atoll (Fig- 
ure 7.19). In the NWHI, eggs are laid 
from December to February, and chicks 
are present from February to May. Most 
birds have dispersed by June. 

The feeding behavior of Tristram's 
Storm-petrels consists of pattering and 
snatching prey from the surface. Gen- 
erally, the diet in the NWHI consists of 
small fish and squid and sometimes 
planktonic halobates (insects) and crus- 
taceans (USFWS, 2005). 

Breeding colonies were extirpated from 
Midway Atoll and Kure Atoll, most likely 
by the introduction of rats. Recoloniza- 
tion is possible due to the eradication 
of rats from these locations by the year 
2000. Nesting habitat degradation from 
invasive plant species continues to be a threat at Pearl and Hermes Atoll. 




Figure 7. 18. Tristram's Storm-petrel, Tern Island and French Frigate Shoals. 
Photo: C. Gregory. 




Figure 7. 19. Tristram's Storm-petrel nesting sites in the NWHI. Source: US- 
FWS, unpub. data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



PELECANIFORMES (BOOBIES, FRIGATEBIRDS AND TROPICBIRDS) 

Red-footed Booby {Sula sula rubripes) 
The Red-footed Booby is the smallest 
of the booby species worldwide, with a 
length of 69-79 cm and body mass of 
850-1,100 g (Schreiberetal.,1996; Fig- 
ure 7.20). Females are slightly larger 
than males. Adults in the NWHI are 
white, often with a yellowish wash on 
the head and neck, with black primaries, 
secondaries and coverts. A tan morph 
also occurs in the NWHI, but is present 
in much lower numbers and is seen only 
rarely (A. Anders, pers. comm.). Bills of 
females are blue with a pink base, and 
those of males lighter blue with a lime 
green and pink base. Legs and feet 
are orange-red to red (Schreiber et al., 
1996). 




Red-footed Boobies are a pantropi- 
cal species, with the largest breeding 
colonies occurring in Palmyra and the 
Hawaiian Islands. Red-footed Boobies 
nest on all of the islands and atolls in the 
NWHI, as well as on Kauai, Oahu and 
offshore islets in the MHI (Figure 7.21). 
The species lays eggs from March to 
August and chicks occur from May to 
December. 

The species forages farther than other 
boobies, ranging up to 276 km from the 
nesting colony (USFWS unpublished 
data; Figure 7.21). During the non- 
breeding period, the birds have been 
observed to travel several hundred kilo- 
meters from land (USFWS, 2005). The 
Red-footed Booby is a plunge diver, with 
primary prey species including flying fish 
and squid. This species often forages in 
large, multi-species flocks associated 
with subsurface predator. 



Figure 7.20. Red-footed Boobies, Tern Island, French Frigate Shoals. Pho- 
to: USFWS. 




Figure 7.21. Red-footed boobies nesting sites and foraging areas in the 
NWHI. Source: USFWS, unpub. data; map: K. Keller. 



Habitat destruction is the main threat to populations in the NWHI (USFWS, 2005). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Masked Booby {Sula dactylatra personata) 
Masked Boobies are the largest of the 
booby species, with a body length of 
74-86 cm, and body mass of 1.2-2.4 
kg. As with many other Pelecaniformes 
species, females are slightly larger than 
males. Adults are white with black-brown 
primaries, secondaries and tail, and pur- 
plish orange feet (Anderson, 1993; Fig- 
ure 7.22). 

Masked Boobies are a pantropical spe- 
cies, with the largest breeding colonies 
occurring on Howland, Baker and Jarvis. 
In the NWHI, the species occurs on all 
of islands and atolls (Figure 7.23). Eggs 
are present from January to July, and 
chicks occur from March to October. 



During the breeding season, birds for- 
age up to 160 km from the breeding col- 
onies (USFWS unpublished data; Fig- 
ure 7.23), and during the non-breeding 
period individuals may travel from 1 ,000 
- 2,000 km from the breeding colonies 
(USFWS, 2005). Masked Boobies are 
plunge divers, with the majority of the 
diet consisting of fish, particularly flying 
fish and jacks. A very small portion of 
the diet is squid (USFWS, 2005). 

The primary threats to the Masked boo- 
by populations in the NWHI are habitat 
destruction and loss of nesting habitat 
due to invasive plants (USFWS, 2005). 




Figure 7.22. Masked Booby with chick, Tern Island, French Frigate Shoals. 
Photo: NOAA. 




Figure 7.23. Masked Booby nesting sites and foraging areas in the NWHI. 
Source: USFWS, unpub. data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Brown Booby {Sula leucogaster plotus) 
The Brown Booby is a medium-sized 
booby, with a body length of 64-85 
cm, a wingspan of 132-155 cm, and 
body mass of 950-1,800 g (Schreiber 
and Norton, 2002). Females are mark- 
edly larger than males. Adults are deep 
brown on the back, head, neck and 
throat, with bright white underparts. In 
the NWHI, adults have a pale yellow bill 
with a light blue base, and bluish yellow 
legs and feet (A. Anders, pers. comm.; 
Schreiber and Norton, 2002; Figure 
7.24). 



The Brown Booby distribution overlaps 
with those of the masked and Red-foot- 
ed Booby species, which are pantropi- 
cal (USFWS, 2005). The largest brown 
booby populations occur in the Hawai- 
ian Islands, including the NWHI and 
MHI. The species nests within all of the 
islands and atolls of the NWHI. Brown 
Boobies lay eggs from February to Au- 
gust, and chicks are present from April 
to October (Figure 7.25). 



Foraging occurs near shore, at 8-70 km 
from land (USFWS, 2005; Figure 7.25). 
The species has been known to travel 
2,000 km from breeding colonies during 
the nonbreeding season, but individuals 
generally remain within 80 km of land 
during the breeding period (USFWS, 
2005). Brown Boobies are plunge div- 
ers, with primary prey species consist- 
ing of flying fish, squid, mackerel scad 
and juvenile goatfish (USFWS, 2005). 

As with the other Booby species, the 
main threat to Brown Booby populations 
in the NWHI is habitat destruction (US- 
FWS, 2005). 




Figure 7.24 
Drury. 



Brown Booby, Tern Island, French Frigate Shoals. Photo: 




Figure 7.25. Brown Booby nesting sites and foraging areas in the NWHI. 
Source: USFWS, 2005; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Great Frigatebird {Fregata minor palmerstoni) 
Frigatebirds have the largest wing-area 
to body mass ratio of any avian species: 
Great Frigatebirds, with a length of 85- 
105 cm and wingspan of 205-230 cm, 
have a body mass of just 1 -1 .8 kg (Gaug- 
er Metz and Schreiber, 2002). Males are 
black or black-brown throughout, with a 
purple/blue/green sheen on the dorsal 
feathers of the neck and nape, and a 
red gular sac. The gular pouch is deflat- 
ed and a pale pink-orange color during 
the non-breeding period, and becomes 
bright red and is inflated during display 
prior to the nesting period. Females are 
significantly larger than males, with a 
white throat, breast, and underparts 
and no (or a greatly reduced) gular sac 
(Gauger Metz and Schreiber, 2002; Fig- 
ure 7.26). 




Figure 7.26. Male Great Frigatebird with inflated gular pouch, Tern Island, 
French Frigate Shoals. Photo: D. Dearborn. 



The distribution of Great Frigatebirds is 
pantropical, with the largest breeding 
populations occurring on Nihoa Island 
and Laysan Island (USFWS, 2005). 
Great Frigatebirds nest on all of the is- 
lands and atolls within the NWHI except 
Gardner Pinnacles and Kure Atoll (Fig- 
ure 7.27). In the NWHI, eggs are laid 
between March and July, and chicks are 
present from April to November. 

Foraging ranges during the breeding 
season have been calculated to be up to 
612 km (Weimerskirch et al., 2004), but 
frigatebirds travel up to 7,000 km dur- 
ing the non-breeding period (Dearborn 
et al., 2003; Figure 7.27). Great Frigate- 
birds cannot swim or land on water, and 
thus use surface dipping and aerial pur- 
suit to capture prey. The main prey spe- 
cies are flying fish and squid (USFWS, 
2005). 




Legend 



Great Frigatebird Nesting Siles 

Great Frigatebird Estimated Foraging Area (612 Km) 



| Papahanaumoku5kea 
Marine National Monument 

2O0 400 600 OD0 

Kilometers 



Figure 7.27. Male Great Frigatebird nesting sites and foraging areas in the 
NWHI. Source: Weimerskirch et al., 2004; map: K. Keller. 



The greatest threats to populations in the NWHI are habitat destruction and disturbance during nesting. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Lesser Frigatebird {Fregata ariel ariel) 
Lesser Frigatebirds are significantly 
smaller than Great Frigatebirds, with 
a mean body length of 75 cm (Lind- 
sey, 1986). Male plumage is similar to 
that of the Great Frigatebird, but with 
overall body plumage of lessers being 
darker black and with a brighter purple 
and green irredescence to the dorsal 
neck and nape feathers (Figure 7.28). 
Lesser males also have a bright white 
line on the underside of the wing, pro- 
viding an obvious distinction in flight to 
the male Great Frigatebird (A. Anders, 
pers. comm.). Female lesser plumage 
is similar to that of female Great Frig- 
atebirds, but with darker black feathers 
dorsally, and a full black hood giving a 
black, rather than white, throat (A. An- 
ders, pers. comm.). 

The pantropical distribution of Lesser 
Frigatebirds lies within that of Great 
Frigatebirds, with the largest breeding 
colonies occurring on Baker and How- 
land Islands (USFWS, 2005). Individu- 
als are observed roosting regularly but 
in low numbers in the NWHI; success- 
ful nesting of a Lesser Frigatebird male 
and Great Frigatebird female (with pro- 
duction of hybrid offspring) has been 
observed in multiple years at French 
Frigate Shoals (Figure 7.29), and one 
pair of Lesser Frigatebirds has been 
documented nesting on Tern Island, al- 
though the nest failed prior to hatching 
(Dearborn and Anders, 2000). 

As with Great Frigatebirds, Lesser Frig- 
atebirds use surface dipping to capture 
prey, and main food items include fly- 
ing fish and squid. Lesser Frigatebirds 




Figure 7.28. Lesser Frigatebird. Photo: USFWS. 




Lesser Frigalebird Nesting Srtes 

J PapahSnaumokuak&a 
Marine National Monument 



Only Tern (stand has been surveyed for Lesser Fngalebirds 



Figure 7.29. Lesser Frigatebird nesting sites in the NWHI. Source: USFWS, 
unpub. data; map: K. Keller. 



travel thousands of kilometers from the breeding colonies during the non-breeding period, but are seen most 
often within 80 km of breeding and roosting islands (USFWS, 2005). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Red-tailed Tropicbird {Phaethon rubricauda melanorhynchos) 
Red-tailed Tropicbirds are a relatively 
small, stout Pelecaniformes species, 
with a body length of 44-47 cm (80-102 
cm including long central tail feathers), 
and body mass of 650-780 g (Schreiber 
and Schreiber, 1993). Plumage is white 
throughout, often with a pale pink wash. 
The bill is red-orange, and a black cres- 
cent eye ring extends to a short eye line 
(Figure 7.30). The tail is short and white, 
with two long red central rectrices. 



The Red-tailed Tropicbird is distributed 
throughout the Indo-Pacific region be- 
tween 35 °N and 30 °S. The largest con- 
centration of breeding birds within the 
Pacific occurs in the NWHI at Midway 
Atoll and Laysan Island (Figure 7.31). 
Other smaller colonies are found on 
all of the other islands and atolls in the 
NWHI, in the MHI, and on other Pacific 
Islands such as Johnston Atoll, Ameri- 
can Samoa, Wake and the Marianas 
(USFWS, 2005). Red-tailed tropicbirds 
lay eggs from March to August and 
chicks occur from April to November. 




Figure 7.30. Red-tailed Tropicbird, Tern Island, French Frigate Shoals. Pho- 
to: C. Gregory. 



Foraging ranges have been calculated 
to be approximately 470 km from the 
breeding colonies, which extends be- 
yond the boundaries of the Monument 
(USFWS unpublished data; Figure 
7.31). Red-tailed Tropicbirds are soli- 
tary, plunge divers and feed mostly on 
flying fish. Other prey species include 
squid, mackerel scad, dolphinfish, trun- 
cated sunfish and balloonfish (USFWS, 
2005). They are not associated with 
subsurface predator schools. 







i * 








Midway 
' '-* Atoll 

Kur» * 

moil 


1 


< 

• 

ri & Hermes 
11 

LiSianskl Laysan 
^■to"L Itltnd 


French 


lokumanarnana 

■ -A Nihca 










R 


■ 








m~ 






Legend 

" Red-tailed Tropicbird Nesting 


Sites 

ed Foraging Areas (468 Km) 
N 

A 

) 

Kilometers 


•\ 




| Red-tailed Tropicbird Estirna 


I PapahanaumokuSkea 
Marine National Monument 

D 200 400 600 ED 












■ 1.. 



In the NWHI, rats on Midway Atoll and Figure 7.31. Red-tailed Tropicbird nesting sites and foraging areas in the 
Kure Atoll, along with invasive plants, NWHI. Source: USFWS, unpub. data; map: K. Keller. 
were the main threats to the species. 

Rats have been eradicated, and USFWS is currently conducting native plant restoration and other alien spe- 
cies removal projects to increase available nesting habitat. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



White-Tailed Tropicbird {Phaethon lepturus dorothea) 
White-tailed Tropicbirds are similar in 
appearance to Red-tailed Tropicbirds, 
but are smaller and more slender, with 
a total length of 60-80 cm (including the 
central rectrices), a wingspan of 90-95 
cm, and a mean body mass of 350 g 
(Lee and Walsch-Mcgehee, 1998). As 
with Red-tailed Tropicbirds, body plum- 
age is white, often with a pink wash, but 
the long central rectrices are white, and 
the bill is more orange than red (Lee and 
Walsch-Mcgehee, 1998; Figure 7.32). 



White-tailed Tropicbirds range through- 
out the tropics with the exception of 
the eastern Pacific and northeastern 
Atlantic. The largest colonies in the 
Pacific occur on American Samoa and 
the MHI. Smaller colonies are found at 
Midway Atoll, Palmyra, Wake and the 
Marianas (USFWS, 2005). Within the 
NWHI, White-tailed Tropicbirds nest 
only at Midway Atoll, where a few pairs 
may breed throughout the year (Figure 
7.33). White-tailed Tropicbirds may be 
competitively excluded from other is- 
lands in the northwestern chain by Red- 
tailed Tropicbirds; in areas of sympatry, 
the larger Red-tailed Tropicbirds do of- 
ten outcompete White-tails for nest sites 
(Harrison, 1990). 




Figure 7.32. 
Starr. 



White-tailed Tropicbird, Sand Island, Midway Atoll. Photo: K. 



White-tailed Tropic birds forage at dis- 
tances up to 120 km from nesting colo- 
nies (USFWS, 2005; Figure 7.33), such 
that the foraging range extends beyond 
the boundaries of the Monument. The 
species are solitary, plunge divers and 
feed mainly on flying fish without aggre- 
gations into multi-species feeding flocks 
with other seabirds. 

Introduced predators are the greatest 
threat throughout the breeding range. 





'? 
















» 












Midway 
Atoll 

• - 


« 










Atoll 


irt & Hermes 
II 

Lisianaki Uysan 




















^ » 




£*. 








- 


. Mokumanamana 

Nihoa 
French ^ «^»* island 














■> 


Leger 

■ 


d 

White-tailed Tropicbird Nesting Sites 
i White-tailed Tropicbird Estimated Foraging Areas (120 Km) 








< 




I 


I N 
| Papahanaumokuakea i 

Marine National Monument f\ 
2DQ 400 600 800 






^■1 









Figure 7.33. White-tailed Tropicbird nesting sites and foraging areas in the 
NWHI. Source: USFWS, unpub. data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

CHARADRIIFORMES (TERNS AND NODDIES) 

Black Noddy {Anous minutus marcusi) 
The Black Noddy is similar in appear- 
ance to the Brown Noddy, but with a 
smaller body size (length 35-40 cm, and 
body mass 85-140 g), and darker brown 
plumage throughout. The head is gray- 
brown, fading to grayish-white on the 
forehead, and with a white crescent on 
the lower eye. The bill is black, and legs 
and feet are reddish brown to orange 
(Gauger, 1999; Figure 7.34). 



Black Noddies are a pantropical spe- 
cies, and the largest colonies in the 
NWHI are at Midway Atoll and Nihoa 
Island (USFWS, 2005; Figure 7.35). In 
the NWHI, Black Noddies can breed 
year around, but in general eggs are laid 
between October and June, and chicks 
are present from December to August. 

Breeding adults forage within 80 km of 
nesting colonies and often forage near 
shore <10 fm (USFWS, 2005; Figure 
7.35). Black Noddies feed by surface 
dipping, and prey species include juve- 
nile and larval goatfish, lizardfish, her- 
ring, flying fish and gobies. 

Habitat loss, predation, invasive spe- 
cies and disturbance are all threats to 
Black Noddy populations. Introduced 
insects at Kure Atoll and increases in 
the invasive golden crown-beard have 
had negative impacts on nesting habitat 
(USFWS, 2005). 




Figure 7.34 
Gregory. 



Black Noddy, Tern Island, French Frigate Shoals. Photo: 




Figure 7.35. Black Noddy nesting sites and foraging areas in the NWHI. 
Source: USFWS, 2005; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Brown Noddy (Anous stolidus pileatus) 
Brown Noddies are a medium-sized 
tern species; they are slightly larger 
than Black Noddies, with a length of 40- 
45 cm and a mean body mass of 180 
g. Body plumage is chocolate-brown 
throughout, with a gray-brown head, 
grayish-white forehead, and white eye 
ring. The bill is black, and legs and feet 
are dark gray to black (Chardine and 
Morris, 1996; Figure 7.36). 

Brown Noddies are a pantropical spe- 
cies, with breeding colonies occurring at 
American Samoa, the Marianas, John- 
ston Atoll and the NWHI. The largest 
colony in the NWHI is on Nihoa Island 
(USFWS, 2005; Figure 7.37). In the 
NWHI, females lay eggs between Janu- 
ary and May, and chicks are present 
from February to November. 

Foraging during the breeding season 
occurs within sight of the nesting colo- 
nies. During the nonbreeding season, 
the birds are known to stay within 100 
km of the nesting colonies (USFWS, 
2005; Figure 7.37). Brown Noddies feed 
by surface dipping, and prey includes 
goatfish, lizardfish, mackerel scad, fly- 
ing fish and squid. (USFWS, 2005) 

Predators are the greatest threat to 
Brown Noddy populations. 




Figure 7.36. 
NOAA. 



Brown Noddy, Tern Island, French Frigate Shoals. Photo: 




Legend 



Brown Noddy Estimated Foraging Areas (100 Km) 

N 
Papahanaumokuakea 
Marino National Monument 

BOO 
> Kilometers 



Figure 7.37. Brown Noddy nesting sites and foraging areas in the NWHI. 
Source: USFWS, 2005; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Blue Noddy of Blue-Gray Noddy {Procelsterna cerulea saxatilis) 
The Blue Noddy is the world's smallest 
tern species. Body plumage is bluish- 
gray throughout, with slightly lighter 
coloration on the head and ventral sur- 
faces, and a white eye ring. Primaries 
and tail are dark gray. Bill and legs dark 
gray, and feet are gray with yellow- 
gray webbing (http://www.state.hi.us/ 
dlnr/dofaw/cwcs/Conservation_need. 
htm#Species; A. Anders, pers. obs.; 
Figure 7.38). 



Blue Noddies are found throughout the 
Pacific, with the largest nesting colonies 
on Nihoa Island and Mokumanamana 
(USFWS, 2005). Blue Noddies nest in 
the eastern portion of the NWHI on Ni- 
hoa Island, Mokumanamana, French 
Frigate Shoals (on La Perouse Pin- 
nacle) and Gardner Pinnacles (Figure 
7.39). On these islands, Blue Noddies 
lay eggs between January and May, 
and chicks are present from January to 
June. There is some variation in breed- 
ing times between islands. The species 
is generally considered a year-round 
resident to the Hawaiian Islands. 




Figure 7.38. Blue Noddy, La Perouse Pinnacle, French Frigate Shoals. 
Photo: D. Wright. 



Blue Noddies feed by surface dipping 
and feed mainly on larval lizardfishes, 
flounders, goatfishes and flying fish. 
They have also been known to take 
squid, crustaceans and halobates (US- 
FWS, 2005). 

Natural predators are the main threat to 
blue noddies. 




- Legend 

■ Blue Noddy Nesllng Sites 

| | Papahanaurnokuakea 

Marine National Monument 



soo 
:] Kilometers 



Figure 7.39. Blue Noddy nesting sites in the NWHI. Source: USFWS, un- 
pub. data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



White Tern {Gygis alba alba) 
White Terns are a medium-sized tern 
species, with a length of 27-33 cm, 
wingspan of 70-87 cm, and body mass 
of 77-157 g. Body plumage is entirely 
white, with a black eye ring. The bill is 
black, and legs and feet are gray-blue 
with yellow-white webs (Niethammer 
and Patrick, 1998; Figures 7.40). 

The White Tern is a pantropical species, 
with the largest colonies in the NWHI 
occurring on Nihoa Island and Midway 
Atoll (Figure 7.41). Other islands includ- 
ing American Samoa and the Marianas 
also support large populations (USFWS, 
2005). In the NWHI the species breeds 
year-around. 

White Terns forage in near-shore waters 
during the breeding season. The spe- 
cies forages by surface diving, plung- 
ing and dipping. Prey includes juvenile 
goatfish and flying fish, while a smaller 
portion of the diet is composed of squid, 
needlefishes, halfbeaks, dolphinfishes 
and blennies (USFWS, 2005). 




Figure 7.40. White 
Photo: NOAA. 



Tern with chick, Tern Island, French Frigate Shoals. 




Figure 7.41. White Tern nesting sites in the NWHI. Source: USFWS, unpub. 
data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Sooty Tern {Sterna fuscata oahuensis) 
The Sooty Tern is a medium-sized tern 
species, with a length of 36-45 cm, a 
wingspan of 82-94 cm, and mean body 
mass of 200 g (Schreiber et al., 2002). 
Body plumage is deep black dorsally 
and on the tail, with a highly-contrast- 
ing white face, throat, and underbelly. 
A white patch on the forehead extends 
to just above the eye, creating a black 
eyeline. The bill, legs, and feet are black 
(Schreiber et al., 2002; Figure 7.42). 

Sooty Terns are a pantropical species. 
The largest colonies in the NWHI occur 
at Laysan Island and Lisianski Island. 
Baker and Jarvis support equally large 
populations (USFWS, 2005; Figure 
7.43). Sooty Terns breed on all islands 
and atolls in the NWHI, with eggs laid 
between March and July and chicks 
present from April to September. 

Sooty Tern foraging distances have 
been calculated to be 740 km (USFWS, 
unpublished data) during the breeding 
season and 5,000 km during nonbreed- 
ing season (Figure 7.43). Sooty Terns 
feed by surface dipping, and main prey 
species include squid, goatfish, fly- 
ing fish and mackerel scad (USFWS, 
2005). 

Introduced predators are a major 
threat to Sooty Terns. While rats have 
been eradicated from the NWHI, cattle 
egrets continue to take chicks at Mid- 
way Atoll (USFWS, 2005). In addition to 
introduced predators, native predators 
such as Great Frigatebirds and Laysan 
Finches also prey on Sooty Tern eggs 
and chicks. 




Figure 7.42. A Sooty Tern at Tern Island, French Frigate Shoals. Photo: C. 
Gregory. 




Figure 7.43. Sooty Tern nesting sites and foraging areas in the NWHI. 
Source: USFWS, unpub. data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Gray-Backed Tern {Sterna lunata) 
Gray-backed Terns are a medium- 
sized Tern, with a length of 35-38 cm 
and body mass of 95-145 g. The upper 
wings, back, and tail are slate-gray, and 
the throat, belly, and underwings are 
white. The head is white with a full black 
cap and black eyeline, and the bill, legs, 
and feet are black (Mostello et al., 2000; 
Figure 7.44). 

Gray-backed Terns are found in the 
tropical and subtropical Pacific, with the 
largest breeding colonies at Lisianski 
Island, Nihoa Island and Laysan Island. 
Smaller colonies are found on John- 
ston, Wake and Jarvis (USFWS, 2005). 
Gray-backed Terns nest on all islands 
and atolls in the NWHI, with eggs laid 
from March to July, and chicks present 
from April to September (Figure 7.45). 

Gray-back Terns have been estimated 
to forage up to 370 km from land, which 
includes areas outside of the Monument 
(USFWS, 2005; Figure 7.45). Gray- 
backed Terns feed by hovering and dip- 
ping; prey species include five-horned 
cowfish, juvenile flying fish, goatfish, 
herring and dolphinfish. Additional prey 
include squid, crustaceans, mollusks, 
and marine and terrestrial insects (US- 
FWS, 2005). 

As with other seabird species, habi- 
tat destruction and disturbance are the 
greatest threats to this species in the 
NWHI (USFWS, 2005). 




Figure 7.44.. Gray-backed Tern. Photo: F. Starr. 









w* 






• V 






J 






* 

Midway 
Anil 

ASH 

LiiMtlskl Laysai 








J 




?'% 


*«i 


ardner 


1 


•i 


TV 




■ 


- 




Reef 


• JJ 


*K* 


lokumanamarw II 
, A Nihoa 11 

* Kblui'i [I 


. 








French 

i i.ijiil^ 
Stioalt 


Legend 




■ Gra) 


'backed Tarn Nesting Sites 

-backed Tern Estimated Foraging Areas (370 Km) 


\ 






: G'ay 




I [Par* 
Mari 

200 


ihSnaLrmokuaikea ^ 
ne National Monument * 

400 600 800 rS 










9 






w m 





Figure 7.45. Gray-backed Tern nesting sites and foraging areas in the 
NWHI. Source: USFWS, unpub. data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Little Tern {Sterna albifrons sinensis) 
The distribution of Little Terns (Figure 
7.46) is pantropical; however, popula- 
tions in the NWHI are very small and 
have been found only at Pearl and 
Hermes and Midway Atolls (USFWS, 
2005), and French Frigate Shoals (A. 
Anders, pers. comm.; Figure 7.47). Lit- 
tle Terns breed in the spring. 

During the breeding season, the spe- 
cies stays within 3 km of the breeding 
colonies while foraging. Little Terns are 
shallow-water plunge divers, with a diet 
consisting of small fish, crustaceans, in- 
sects, annelids and mollusks (USFWS, 
2005). 

Habitat destruction and disturbance are 
two of the threats faced by Little Terns. 
The breeding-season foraging range is 
completely protected within the bound- 
aries of the Monument. 




Figure 7.46. Little Tern. Photo: D. Mason, www.realbirder.com. 




Figure 7.47. Little Tern nesting sites in the NWHI. Source: USFWS, unpub. 
data; map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



POPULATION STATUS AND TRENDS 



Available Data 

Large-scale monitoring of seabird populations in the NWHI began in the early 1960s, with the Smithsonian 
Institution's Pacific Ocean Biological Survey Program (POBSP). This program was followed by Tripartite Co- 
operative Agreement Studies in the late 1970s and early 1980s, and by the current long-term monitoring pro- 
gram initiated in 1980 by the USFWS at French Frigate Shoals, Midway Atoll and Laysan Island. The current 
USFWS seabird monitoring program provides information on seabird populations over time within French Frig- 
ate Shoals, Midway and Laysan, but geographical comparisons are difficult, as data collection methods have 
differed across these locations. 



Pacific Ocean Biological Survey Program 

The POBSP was conducted throughout the Pacific by the Smithsonian Institution from 1 963-1 970. The Smith- 
sonian's surveys included at-sea bird observations, as well as breeding population counts and banding on 
islands and atolls within the NWHI from 1 963-1 968 (King, 1 974). The resulting data provide a general baseline 
for five species of seabirds that breed on Nihoa, Mokumanamana, French Frigate Shoals, Lisianski, Laysan, 
Pearl and Hermes Atoll, Midway Atoll and Kure Atoll, including Laysan and Black-footed Albatrosses, Wedge- 
tailed Shearwaters, Red-tailed Tropicbirds and Sooty Terns (Table 7.4). Because the POBSP breeding popu- 
lation size data presented in King (1974) were not well documented in terms of the types and timing of the 
counts, it is not entirely clear if population estimates were total number of individuals or number of breeding 
pairs. It is thus difficult to compare the 1960s data to current population size estimates. However, the POBSP 
data do allow for general comparisons of population sizes between islands and atolls for the time period during 
which those data were collected (Table 7.5). 



Table 7.4. Smithsonian Institution Pacific Ocean Biological Survey Program Survey Sites. 





BLACK-FOOTED LAYSAN'S ffiopiCBIRD SOOTY TERN ^IWATER 
ALBATROSS ALBATROSS Tp h «,hn„ mhri«,,,w a (Sterna fuscata f p H ,l* R c „*!! " c 

(Phoebastrianigripes) (Phoebastria immutabilis) <J^SoS ) rubncauda oahuensis) S£»? 


French Frigate 
Shoals 


X 


X 


X 


X 


X 


Kure 


X 


X 


X 


X 


X 


Laysan 


X 


X 


X 


X 


X 


Lisianski 


X 


X 


X 


X 


X 


Midway 


X 


X 


X 


X 


X 


Mokumanamana 


X 


X 


X 


X 


X 


Nihoa 


X 


X 


X 


X 


X 


Pearl and Hermes 


X 


X 


X 


X 


X 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 7.5. 


POBSP Seabird Population Survey Results. 












SITE 


SPECIES 


SCIENTIFIC NAME 


COMMON 
NAME 


DATE 


LOW 
VALUE 


V 


Hpf 




Lisianski 


LAAL 


Phoebastria immutabilis 


Laysan Albatross 


1963-1968 


8,000 


10,000 


Largest number of breeders 


Nihoa 


LAAL 


Phoebastria immutabilis 


Laysan Albatross 


1963-1968 


- 


38 


Largest number of breeders 


Kure 


LAAL 


Phoebastria immutabilis 


Laysan Albatross 


1963-1968 


- 


3,200 


Largest number of breeders 


MMM 


LAAL 


Phoebastria immutabilis 


Laysan Albatross 


1963-1968 


- 


1,650 


Largest number of breeders 


FFS 


LAAL 


Phoebastria immutabilis 


Laysan Albatross 


1963-1968 


- 


1,000 


Largest number of breeders 


Midway 


LAAL 


Phoebastria immutabilis 


Laysan Albatross 


1963-1968 


- 


110,000 


Largest number of breeders 


Laysan 


LAAL 


Phoebastria immutabilis 


Laysan Albatross 


1963-1968 


300,000 


500,000 


Largest number of breeders 


Pearl and 
Hermes 


LAAL 


Phoebastria immutabilis 


Laysan Albatross 


1963-1968 


- 


30,000 


Largest number of breeders 


MMM 


BFAL 


Phoebastria nigripes 


Black-footed 
Albatross 


1963-1968 


- 


375 


Maximum number of breed- 
ing birds recorded 


FFS 


BFAL 


Phoebastria nigripes 


Black-footed 
Albatross 


1963-1968 


- 


3,100 


Maximum number of breed- 
ing birds recorded 


Laysan 


BFAL 


Phoebastria nigripes 


Black-footed 
Albatross 


1963-1968 


- 


40,000 


Maximum number of breed- 
ing birds recorded 


Nihoa 


BFAL 


Phoebastria nigripes 


Black-footed 
Albatross 


1963-1968 


- 


100 


Maximum number of breed- 
ing birds recorded 


Lisianski 


BFAL 


Phoebastria nigripes 


Black-footed 
Albatross 


1963-1968 


- 


4,000 


Maximum number of breed- 
ing birds recorded 


Pearl and 
Hermes 


BFAL 


Phoebastria nigripes 


Black-footed 
Albatross 


1963-1968 


- 


9,000 


Maximum number of breed- 
ing birds recorded 


Midway 


BFAL 


Phoebastria nigripes 


Black-footed 
Albatross 


1963-1968 


- 


24,000 


Maximum number of breed- 
ing birds recorded 


Kure 


BFAL 


Phoebastria nigripes 


Black-footed 
Albatross 


1963-1968 


- 


1,150 


Maximum number of breed- 
ing birds recorded 


MMM 


WTSH 


Puffinus pacificus 
chlororhynchus 


Wedge-tailed 
Shearwater 


1963-1968 


- 


2,000 


Maximum number recorded 


Kure 


WTSH 


Puffinus pacificus 
chlororhynchus 


Wedge-tailed 
Shearwater 


1963-1968 


- 


6,230 


Maximum number recorded 


MMM 


SOTE 


Sterna fuscata 
oahuensis 


Sooty Tern 


1968 


- 


50,000 


Maximum number recorded 


Midway 


SOTE 


Sterna fuscata 
oahuensis 


Sooty Tern 


1968 


- 


353,000 


Maximum number recorded 


Laysan 


SOTE 


Sterna fuscata 
oahuensis 


Sooty Tern 


1968 


- 


2,000,000 


Maximum number recorded 


Nihoa 


SOTE 


Sterna fuscata 
oahuensis 


Sooty Tern 


1968 


- 


100,000 


Maximum number recorded 


Pearl and 
Hermes 


WTSH 


Puffinus pacificus 
chlororhynchus 


Wedge-tailed 
Shearwater 


1963-1968 


- 


22,400 


Maximum number recorded 


Laysan 


WTSH 


Puffinus pacificus 
chlororhynchus 


Wedge-tailed 
Shearwater 


1963-1968 


- 


200,000 


Maximum number recorded 


FFS 


SOTE 


Sterna fuscata 
oahuensis 


Sooty Tern 


1968 


- 


250,000 


Maximum number recorded 


Pearl and 
Hermes 


SOTE 


Sterna fuscata 
oahuensis 


Sooty Tern 


1968 


- 


40,110 


Maximum number recorded 


Nihoa 


WTSH 


Puffinus pacificus 
chlororhynchus 


Wedge-tailed 
Shearwater 


1963-1968 


- 


25,000 


Maximum number recorded 


Kure 


SOTE 


Sterna fuscata oahuen- 
sis 


Sooty Tern 


1968 


- 


48,000 


Maximum number recorded 


Lisianski 


WTSH 


Puffinus pacificus 
chlororhynchus 


Wedge-tailed 
Shearwater 


1963-1968 


- 


60,000 


Maximum number recorded 


Lisianski 


SOTE 


Sterna fuscata 
oahuensis 


Sooty Tern 


1968 


- 


1,000,000 


Maximum number recorded 


Midway 


WTSH 


Puffinus pacificus 
chlororhynchus 


Wedge-tailed 
Shearwater 


1963-1968 


- 


3,000 


Maximum number recorded 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 7.5. (continued). POBSP Seabird Population Survey Results. 



SPECIES SCIENTIFIC NAME 



Nihoa 
Kure 



RTTR 
RTTR 



COMMON 
NAME 



Phaethon rubricauda 
rothschildi 

Phaethon rubricauda 
rothschildi 



Red-tailed 
Tropicbird 

Red-tailed 
Tropicbird 



1963-1968 
1963-1968 



HIGH 
VALUE 



500 
2,500 






Maximum population estimates 
Maximum population estimates 



MMM RTTR 

FFS RTTR 

Laysan RTTR 



Phaethon rubricauda 
rothschildi 

Phaethon rubricauda 
rothschildi 

Phaethon rubricauda 
rothschildi 



Red-tailed 
Tropicbird 

Red-tailed 
Tropicbird 

Red-tailed 
Tropicbird 



1963-1968 
1963-1968 
1963-1968 



200 

225 

4,000 



Maximum population estimates 
Maximum population estimates 
Maximum population estimates 



Pearl and RTTR 
Hermes 



Phaethon rubricauda 
rothschildi 



Red-tailed 
Tropicbird 



1963-1968 



165 



Maximum population estimates 



Lisianski RTTR 



Midway RTTR 



Phaethon rubricauda 
rothschildi 



Red-tailed 
Tropicbird 



Phaethon rubricauda 
rothschildi 



1963-1968 



1963-1968 



Red-tailed 
Tropicbird 

Island/atoll abbreviations: MMM = Mokumanamana, FFS = French Frigate Shoals 

Species abbreviations: LAAL = Laysan's Albatross; BFAL = Black-footed Albatross; WTSH 
Tern; WTSH = Wedge-tailed Shearwater; RTTR = Red-tailed Tropicbird 



3,000 
7,500 



Maximum population estimates 
Maximum population estimates 



Wedge-tailed Shearwater; SOTE = Sooty 



Tripartite Cooperative Agreement Studies 

In 1975, the National Marine Fisheries Service, USFWS, and Hawaii Division of Aquatic Resources (then 
known as Division of Fish and Game) established a five-year Tripartite Cooperative Agreement to conduct sur- 
veys and assessments of the NWHI (Grigg and Tanoue, 1 984). One of the publications resulting from this co- 
operative agreement included a summary of the seabird research that had been conducted in the northwestern 
islands, and included information on 18 species of seabirds that breed in the northwestern chain. From 1978 
to 1982, USFWS conducted field studies of variable length (from five weeks to year-around) at Nihoa Island, 
French Frigate Shoals, Laysan Island and Lisianski Island. In addition, short field trips were conducted to most 
of the other islands and atolls, but these shorter trips did not produce detailed population evaluations. The data 
presented in the Tripartite Cooperative Agreement Studies report is a combination of POBSP data and the field 
data collected from 1978-1982. The report is limited in use for biogeographic comparisons between islands/ 
atolls because the dates and methods of data collection were not reported, but it is possible to use the data as 
a baseline for seabird breeding population sizes within islands and atolls (Fefer et al., 1984). 

USFWS Seabird Monitoring Program 

The USFWS has been conducting seabird monitoring at French Frigate Shoals and Midway Atoll since 1980. 
Various levels of population monitoring have been conducted at French Frigate Shoals for each of the fol- 
lowing species: Laysan Albatrosses, Black-footed Albatrosses, Bonin Petrels, Bulwer's Petrels, Wedge-tailed 
Shearwaters, Christmas Shearwaters, Tristram's Storm-petrels, Red-footed Boobies, Masked Boobies, Great 
Frigatebirds, Red-tailed Tropicbirds, Black Noddies, Brown Noddies, White Terns and Gray-backed Terns. 
Monitoring at Laysan Island also occurs once a year, where data are collected on Laysan Albatrosses, Black- 
footed Albatrosses, Red-footed Boobies and Great Frigatebirds (data from 1992-present are currently avail- 
able). As time and funding allow, periodic monitoring also occurs on the other islands and atolls in the NWHI. 
The long-term data sets from French Frigate Shoals, Midway Atoll and Laysan Island provide information 
about the variability of the seabird populations within each of those islands/atolls over the past 27 years, but 
because monitoring methods have differed between locations, inter-island or-atoll comparisons for biogeo- 
graphic assessment are limited. As a first step toward future biogeographic analysis, data layers of presence 
and absence of species are presented here, as are a subset of results from the long-term seabird monitoring 
program at French Frigate Shoals and Midway Atoll. Table 7.5 also indicates the types of monitoring data that 
have been collected at each island and atoll. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

USFWS Monitoring at French Frigate Shoals 

Methods 

Since 1 980, the USFWS has conducted field monitoring on Tern Island, French Frigate Shoals, to estimate the 
minimum number of breeding pairs and annual reproductive success of a subset of the seabird species that 
breed on the island. Beginning in 1980-1981, minimum numbers of breeding pairs and reproductive success 
were estimated annually for Laysan and Black-footed Albatrosses, Red-footed Boobies, Red-tailed Tropic- 
birds, Black Noddies, and White Terns (Dearborn et al., 2001). USFWS has since expanded the monitoring 
program on Tern Island, such that minimum numbers of breeding pairs are now estimated annually, as of 
2008, for Laysan Albatrosses, Black-footed Albatrosses, Christmas Shearwaters, Bulwer's Petrels, Tristram's 
Storm-petrels, Red-footed Boobies, Masked Boobies, Great Frigatebirds, Red-tailed Tropicbirds, Black Nod- 
dies, Brown Noddies, White Terns and Gray-backed Terns (USFWS, 2008). In addition, annual reproductive 
success is now monitored for Laysan and Black-footed Albatrosses, Bulwer's Petrels, Tristram's Storm-petrels, 
Red-footed Boobies, Masked Boobies, Red-tailed Tropicbirds and Black Noddies (USFWS, 2008). 

Estimation of the minimum number of pairs breeding on Tern Island varies by species, depending primarily 
upon population size and the level of breeding synchrony exhibited by the species. For the Procellariiformes, 
including Laysan and Black-footed Albatrosses, Christmas Shearwaters, Bulwer's Petrels, and Tristram's 
Storm-petrels, population sizes are relatively small, and breeding synchrony is high. For these reasons, the 
number of breeding pairs nesting on Tern Island are counted directly each year for each of these species 
(numbers of breeding pairs of Laysan and Black-footed Albatrosses are also directly counted each year at all 
other islands within French Frigate Shoals, such that the number of breeding pairs for the entire atoll can be 
estimated for these two species). In contrast, breeding within the Pelecaniformes and Charadriiformes spe- 
cies is highly asynchronous, and population sizes are larger, such that direct counts of breeding pairs are not 
possible (with the exception of Masked Boobies, for which all breeding adults on Tern Island are banded, and 
the breeding population consists of only a few hundred pairs; A. Anders, pers. comm.). For these species, es- 
timates of the minimum numbers of breeding pairs are based upon mean incubation count nest censuses: for 
each species, a count of all nests at which an adult is incubating is conducted at a periodicity that equals the 
mean incubation length for that species. For example, the mean incubation length of Great Figatebirds is 55 
days, so a count of all nests at which an egg is being incubated is conducted every 55 days (USFWS, 2008). 
Summing the mean incubation counts over an entire year then provides information on the minimum number of 
breeding pairs that attempted to nest in that year (the estimate is a minimum, as some nests are initiated and 
fail between mean incubation count periods, so these failed breeding pairs are not counted; there is also some 
error introduced by the fact that within some species, individuals or breeding pairs may attempt to re-nest after 
nest failure within a year). 

Finally, annual reproductive success is also monitored based upon species' population sizes and level of 
breeding synchrony. For Laysan Albatrosses, Black-footed Albatrosses, Bulwer's Petrels, Tristram's Storm- 
petrels, Masked Boobies, White Terns, and Gray-backed Terns, breeding populations are small enough and/or 
breeding is synchronous enough that virtually all nests on Tern Island are monitored to obtain an annual esti- 
mate of nesting success. In contrast, for Red-footed Boobies, Red-tailed Tropicbirds, and Black Noddies, nests 
are monitored year-around within multiple randomly-chosen permanent plots on Tern Island, with all nests in all 
plots being checked every two to four days for failure, hatching, or fledging (Dearborn and Anders, 1996). For 
each of these species, the reproductive success plots include approximately 20-30% of all nests on Tern Island 
for that particular species (Dearborn et al., 2001). Randomly-chosen permanent nest monitoring plots were 
also set up for Laysan and Black-footed Albatrosses beginning in 2006, in order to obtain more precise esti- 
mates of nesting success on Tern Island for these two species (USFWS, 2008; A. Anders, pers. comm.). For 
all species, nest monitoring methods provide data for annual estimates of hatching success, fledging success, 
and overall reproductive success for each species monitored. Estimates of minimum numbers of breeding 
pairs and for hatching, fledging, and overall reproductive success are available for all monitored species from 
1996 through the present from the Pacific Seabird Monitoring Database (Pacific Seabird Group and USGS, 
Alaska Biological Science Center, 2007). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Results 

The following figures (Figure 7.48) in- 
dicate the minimum numbers of breed- 
ing pairs of Laysan Albatrosses and 
Black-footed Albatrosses each year at 
all islands within French Frigate Shoals 
from 1 960 - 2007 (1 962 data: Rice and 
Kenyon, 1962; 1960-1983 data: Harri- 
son et al., 1983; 1980-2007 data: Pa- 
cific Seabird Group and USGS, 2007). 
For both species, breeding population 
size varies annually, but both species 
have exhibited population increases at 
French Frigate Shoals since 2004. 

Estimates of the minimum numbers of 
breeding pairs on Tern Island, French 
Frigate Shoals are presented in Fig- 
ure 7.49 for Bulwer's Petrels, Christ- 
mas Shearwaters, Red-footed Boobies, 
Masked Boobies, Red-tailed Tropic- 
birds, Great Frigatebirds, Black Nod- 
dies, Brown Noddies, White Terns and 
Gray-backed Terns from 1996 - 2006 
(Pacific Seabird Monitoring Database, 
2008). In 2006, Bulwer's Petrel nest 
boxes were temporarily relocated dur- 
ing building construction on Tern Island, 
such that only natural crevices were 
available for nesting for this species 
in that year. The number of Christmas 
Shearwater breeding pairs appears to 
have declined in 2002-2006 relative to 
steadily over the 11 -year period shown, 
or remained stable from 1996-2006. 



4500 -I 

4000 Laysan Albatross 

3500 - 
3000 - 
2500 - 
2000 - 
1500 - 
1000 - 
500 - 
— 



& j8> j8> jfr j8> 



& _<# _(& .<# J» „oP _o> J* „<S> J& J? .<*> JS Jt* J# J* J^ -c?- -C?> J* JP -sA 



&^&&&&&&&&&&&&&&&&fw&f^tfty'J r #'# 



Black-footed Albatross 



.± 5000 - 

re 

Q. 



o> os° o, s o!" o, k ^= oS 1 oJ o!> o? tS> o> dJ- (v> a 1, (S o? dj a* ^ t? I\ N * c? 1 R k l^> 1^ 

N C?> n O? ^> N # N # N # N # N C?> n O?> N C?> N C? n O? n O? N C? s O? N # N C? ^ N of» n # rj> cf cj> r£ Jj Q Q ^ 

? 

Hatch Year 



Figure 7.48. The minimum numbers of breeding pairs of Laysan Albatross- 
es and Black-footed Albatrosses each year at all islands within French Frig- 
ate Shoals from 1960 -2007. Sources: Rice and Kenyon, 1962; Harrison, 
1983; Pacific Seabird Monitoring Database, 2008. 

1996-2001, and numbers of Masked Booby breeding pairs increased 
All other species' breeding population sizes have fluctuated annually 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



100 
90 
80 
70 
60 
50 
40 
30 
20 
10 



Bulwer's Petrel 



H 



f 



<2 3000 

Q. 

Dl 2500 



m 



2000 

1500 
1000 
500 



1200 
11000 
800 
600 
400 
200 



Red-footed Booby 



</ ^ 






Red-tailed Tropicbird 



6000 
' 5000 
4000 
3000 
2000 
1000 



.t& _cj jd?> j& 



Black Noddy 



<* c?> q?> o? 

■f s# rf S* 



s? o N & c?> a 1 * <&> 6 



# # 



/ ^ 



Christmas Shearwater 



# ^ 



/ ^ 



160 
140 
120 
100 



Masked Booby 



# ,# A # .# ^ „# „# ^ / J= rf 



soo -j Great Frigatebird 

700 - 
600 - 
500 - 
400 - 
300 - 
200 - 
100 - 



•f f f <F f V V 



<* £■ r& 



m 6000 

'5 

a. 

D) 5000 

B 

T3 

| 4000 

CD 

° 3000 
3 

-Q 

3 2000 
Z 

E 

=> 1000 

E 



rit» rW rsv «J r*J r*-> ri^ r4>J 

■9 <?> ■? ^ i? i? <£> <£> 



Brown Noddy 



<# <$ <# <# # c? N # C# c# # c? fc 

>J> ,°> ,°> k°> „9> o „s> „9> fP oV o? 



Gray-backed Tern 



# # 



/ / 



Figure 7.49. Estimates of the minimum numbers of breeding pairs on Tern Island, French Frigate Shoals from 1996-2006. 
Source: Pacific Seabird Monitoring Database, 2007. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Reproductive Success 

Annual nesting success (total number of chicks fledged/total number of eggs laid) from 1996 - 2006 is pre- 
sented in Figure 7.50 below for Laysan and Black-footed Albatrosses, Bulwer's Petrels, Red-footed Boobies, 
Masked Boobies, Red-tailed Tropicbirds, Black Noddies, and Gray-backed Terns. Black-footed Albatross re- 
productive success has remained relatively stable - between approximately 70-80% - over the 11 -year period, 
while Laysan Albatross success has been lower and more variable - between 30% and just over 70% - during 
the same time period. Reproductive success for Red-tailed Tropicbirds and Gray-backed Terns appears to 
have declined substantially after the 1 996 and 1 997 breeding seasons, while all other species' nesting success 
fluctuated annually over that time period. Cause of the extremely low reproductive success for Black Noddies 
in 2002 is unknown, although for all species, overall nesting success is tied very closely to food availability in 
any given year (2002-2003 was an El Nino year, such that food availability may have been low for some sea- 
bird species during that time). 



£100 
in 90 



§ 60 
? 50 
| 40 

1 30 
!E 20 

2 10 



o 50 



CO 40 



o 30 



DC 
= 10 



> 
O 



«i 60 
n 

o> 

u 50 

en 

| 40 

o 

•d 30 
o 

a. 

£ 20 

~m 

5 10 
> 
O 




Black-footed Albatross 



1996 1997 1998 2000 2001 2002 2003 2004 2005 2006 



Bulwer's Petrel 



1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 



-? -. Masked Booby 



in 70 

o 

u 

^ 60 

CO 

o 50 

> 

o 40 



20 
10 - 



1996 1997 1998 1S 



Black Noddy 



2000 2001 2002 2003 2004 2005 2006 



1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 



£ 80 

tn 70 

Q 

U 

60 

3 

"> „ 

a, 50 

> 

'■5 40 

| 30 

1 20 

B 10 



25 



3 20 



1 10 



Laysan Albatross 



1996 1997 1998 2000 2001 2002 2003 2004 2005 2006 



£ 60 

l 50 

o 

o 

S> 40 
0) 

> 

5 30 

3 

■o 
o 

£ 20 
a> 

X 

= 10 
re 

q 

~ 60 



Red-footed Booby 



1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 



Red-tailed Tropicbird 



40 



30 



20 



B 10 



40 
f 35 

01 


K 30 



fl 



1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 



Grav-backed Tern 



n 



1996 1997 18 



2000 2001 2002 2003 2004 2005 



Figure 7.50. Annual nesting success (total number of chicks fledged/total number of eggs laid) from 1996 - 2006. Source: 
Pacific Seabird Monitoring Database, 2007. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



USFWS Monitoring at Midway Atoll 

Methods 

Laysan and Black-footed Albatross 
nests have been counted at Midway 
Atoll since 1 991 , and beginning in 1 997, 
reproductive success of both species 
has been monitored. Hatching success, 
fledging success, and overall reproduc- 
tive success, have been estimated for 
both species from 2002 - 2008 (Klavit- 
ter et al., 2009). Red-tailed Tropicbird 
population parameters have also been 
studied at Midway Atoll since 1997, 
including reproductive success, adult 
survival, and overall population trends 
(Laniawe and Klavitter, 2009; Laniawe, 
2008). Seabird monitoring at Mid- 
way also currently includes population 
monitoring of Short-tailed Albatrosses, 
Brown Boobies, Masked Boobies, Bul- 
wer's Petrels, Tristram's Storm-petrels, 
Least Terns and Little Terns (J. Klavitter, 
pers. comm.). 

Results 

Results of Laysan and Black-footed Al- 
batross nest monitoring from 2002-2008 
are presented in Figure 7.51. For this 
seven-year period, mean Laysan Alba- 
tross hatching success was 82% and 
fledging success was 85%, leading to 
an overall reproductive success of 69%. 
Black-footed Albatrosses had similar 
population paramaters, with a hatching 
success of 83%, fledging success of 
83%, and overall reproductive success 
of 70% for the entire time period, In 
analyzing reproductive parameters be- 
tween species and years, Black-footed 
Albatrosses had a slightly higher hatch- 
ing success than Laysan Albatrosses, 
but the two species had similar fledging 
and overall reproductive success rates 
(Klavitter et al., 2009). 



Laysan Albatross 

□ Hatching □ Fledging □ Reproductive 

1.00 
0.90 
0.80 
0.70 
0.60 
0.50 
0.40 
0.30 
0.20 
0.10 
0.00 







JL 


























JL. 










JL. 


JL 


jl 














JL 








JL 






JL 








JL 








_1_ 




















































JL 








JL 




























































- 

































































































































































































































































































2002 2003 2004 2005 2006 2007 2008 



Black-footed Albatross 



□ Hatching □ Fledging □ Reproductive 



1.00 
0.90 
0.80 
0.70 
in 0.60 
H 0.50 



(ft 



0.40 
0.30 
0.20 
0.10 
0.00 













_L 


JL 


iX 


JL 












JL 




J_ 














X 












JL 








_L 






JL 




J_ 


























_L 
























LL 








JL 

































































































































































































































































































































































2002 2003 2004 2005 2006 2007 2008 



Figure 7.51. The top panel shows Laysan Albatross hatching, fledging, and 
overall reproductive success at Midway Atoll from 2002-2008. The bottom 
panel Black-footed Albatross hatching, fledging, and overall reproductive 
success at Midway Atoll from 2002-2008. Source: Klavitter et al., 2009. 



Red-Tailed Tropicbird 

□ Hatching □ Fledging □ Reproductive □ Survivorship 


.9- 0.80 - 


T 




L 

Tt t 


T * 




I _JL_ 


■- I 


I 






r J 


i 

T 




U 


Tt 


I 


T[ 




3 
(f) 


T 


T 




I 




1 


I 


rM. 


1 


i 




</> 


1 




I 


I 




T 


I 








T 


O 

u 


I 








I 






1 


I 


1 


1 


O) 
























CO 

0) 
















































1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 



The results of Red-tailed Tropicbird 
population monitoring at Midway Atoll 
from 1997-2007 are presented in Fig- 
ure 7.52. Hatching, fledging, and overall 
reproductive success, as well as adult 
survival, varied between years, but for 

the entire 1 1 -year period, reproductive success was 41 %, and adult survival was 70%. These reproductive and 
survival rates led to an overall population trend of 0.82 for this time period, indicating that the Red-tailed Trop- 
icbird population at Midway Atoll was declining during this time (Laniawe and Klavitter, 2009; Laniawe, 2008). 



Figure 7.52. Red-tailed Tropicbird reproductive success and survival es- 
timates for Midway Atoll from 1997-2007. Source: Laniawe and Klavitter, 
2009. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

EXISTING DATA GAPS 

Spatial Distribution 

Limiting the population monitoring data to three of the islands/atolls within the NWHI reduces our ability to 
analyze population productivity, survival and community composition across the archipelago. The remoteness, 
year-around breeding, and underground nesting habits of some of the petrel species make breeding seabird 
surveys at all of the islands within the Monument logistically difficult. Nest density calculations at the various 
islands, along with information regarding available habitat (and the extent covered by introduced plants), might 
allow for analysis and predictions of future impacts from changing sea levels and vegetation change over time. 
In addition, standardized methods for monitoring reproductive success and survival at French Frigate Shoals, 
Laysan and Midway Atoll would allow for more comprehensive geographic comparisons of seabirds nesting 
within the Monument. The collection of at sea seabird survey data should be conducted on a regular basis in 
order to gain a better understanding of the at sea behavior and environmental conditions for foraging. Currently 
a single survey was conducted by the NOAA Pacific Southwest Fisheries Science Center in 2002. This type of 
survey needs to be conducted again and at different times of year in order to gain a better understanding of all 
the seabirds in the NWHI throughout the year. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

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Harrison, OS. 1990. Seabirds of Hawaii: Natural History and Conservation. Cornell University Press, Ithaca, New York. 

Harrison, OS., T.S. Hida, and M.P. Seki. 1983. Hawaiian seabird feeding ecology. Wildlife Monographs 85: 3-71 . 

Harrison, P. 1987. A Field Guide to Seabirds of the World. The Stephen Greene Press, Lexington, Massachusetts. 

Hyrenbach, K.D., P. Fernandez and D.J. Anderson. 2002. Oceanographic habitats of two sympatric North Pacific alba- 
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Hyrenbach, K.D., Keiper, O, Allen, S.G., Ainley, D.G., and D.J. Anderson. 2006. Use of marine sanctuaries by far-ranging 
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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Kappes, M.A., S.A. Shaffer, Y. Tremblay, D.G. Foley, D.M. Palacios, P.W. Robinson, S.J. Bograd and D.P. Costa, (in 
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Klavitter, J., Schubert, G., Laniawe, L, and M. Romano. 2009. Reproductive success of Laysan and black-footed alba- 
trosses from 2002 - 2008 at Midway Atoll National Wildlife Refuge. Pacific Seabird Group Annual Meeting. 1 8 - 26 Febru- 
ary, 2009. Hakodate, Hakkaido, Japan. 

Kurata, Y. 1 978. Breeding record of the Laysan Albatross Diomedea immutabilis on the Ogasawara Islands, Japan. Jour- 
nal of the Yamashina Institute for Ornithology 1 0: 1 85-1 89. 

Laniawe, L. 2008. Survivorship, Productivity, and Foraging Range of the Red-tailed Tropicbird (Phaethon rubricauda) at 
Midway Atoll National Wildlife Refuge. Thesis. University of East Anglia, Norwich, England, UK. 

Laniawe, L. and J. Klavitter. 2009. Reproductive success, survivorship, and population trend of the red-tailed tropicbird 
1997 - 2007 at Midway Atoll National Wildlife Refuge. Pacific Seabird Group Annual Meeting. 18-26 February, 2009. 
Hakodate, Hakkaido, Japan. 

Lee, D.S. and M. Walsh-Mcgehee. 1998. White-tailed Tropicbird (Phaethon lepturus), The Birds of North America Online 
(A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna. birds. 
cornell.edu/bna/species/353. 

Lewison, R. and L.B. Crowder. 2003. Estimating fishery bycatch and effects on a vulnerable seabird population. Ecologi- 
cal Applications 13 (3): 743-753. 

Lindsey, T.R. 1 986. The seabirds of Australia. Angus and Robertson, Australia. 

Megysi, J.L. and D.L. O'Daniel. 1997. Bulwer's Petrel (Bulweha bulwerii), The Birds of North America Online (A. Poole, 
Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/ 
bna/species/281 . 

Mostello, C.S., N.A. Palaia and R.B. Clapp. 2000. Gray-backed Tern (Sterna lunata), The Birds of North America Online 
(A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna. birds. 
cornell.edu/bna/species/525. 

Niethammer, K.R. and L.B. Patrick. 1998. White Tern (Gygis alba), The Birds of North America Online (A. Poole, Ed.). 
Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/ 
species/371. 

Pacific Seabird Group and USGS, Alaska Biological Science Center, 2007 Pacific Seabird Monitroing Database, http:// 
www.absc.usgs.gov/research/psinfonet/psmdb/splashpsmdb.htm 

Rice, D. W. and K. W. Kenyon. 1962. Breeding distribution, history and populations of North Pacific Albatrosses. Auk 79: 
365-386. 

Schreiber, E.A., C.J. Feare, B.A. Harrington, B.G. Murray, Jr., W.B. Robertson, Jr., M.J. Robertson and G.E. Woolfenden. 
2002. Sooty Tern (Sterna fuscata), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; 
Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/species/665 

Schreiber, E.A. and R.L. Norton. 2002. Brown Booby (Sula leucogaster), The Birds of North America Online (A. Poole, 
Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/ 
bna/species/649 

Schreiber, E.A. and R.W. Schreiber. 1993. Red-tailed Tropicbird (Phaethon rubricauda), The Birds of North America On- 
line (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http ://bna. birds. 
cornell.edu/bna/species/043 

Schreiber, E.A., R.W. Schreiber and G.A. Schenk. 1996. Red-footed Booby (Sula sula), The Birds of North America On- 
line (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http ://bna. birds. 
cornell.edu/bna/species/241 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Seto, N.W. 2001. Christmas Shearwater (Puffinus nativitatis) , The Birds of North America Online (A. Poole, Ed.). 
Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/ 
species/561 

Seto, N.W. and D. O'Daniel. 1999. Bonin Petrel (Pterodroma hypoleuca), The Birds of North America Online (A. Poole, 
Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/ 
bna/species/385 

Shaffer, S.A., D.M. Palacios, Y. Tremblay, M.A. Kappes, D.G. Foley, S.J. Bograd, and D.P Costa, (in review). A tale of two 
hotspots. Segregation at sea by post-breeding Hawaiian albatrosses. Journal of Animal Ecology. 

Shaffer, S.A., Y. Tremblay, J.A. Awkerman, R.W. Henry, S.L.H. Teo, D.J. Anderson, D.A. Croll, B.A. Block and D.P. Costa. 
2005. Comparison of light- and SST-based geolocation with satellite telemetry in free-ranging albatrosses. Marine Biology 
147:833-843 

Slotterback, J. W. 2002. Tristram's Storm-petrel (Oceanodroma tristrami), The Birds of North America Online (A. Poole, 
Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/ 
bna/species/673b 

Suryan, R.M., D.J. Anderson, S.A. Shaffer, D.D. Roby, Y Tremblay, D.P. Costa, P.M. Sieved, F. Sato, K. Ozaki, G.R. Ba- 
logh and N. Nakamura.(in press) Wind, waves, and wing-loading: Specialization in two Pacific albatross species. PLoS 
One. 

Suryan, R.M., F. Sato, G.R. Balogh, K.D. Hyrenbach, P.R. Sievert, and K. Ozaki. 2006. Foraging destinations and marine 
habitat use of short-tailed albatrosses: A multi-scale approach using first-passage time analysis. Deep-Sea Research Part 
li-Topical Studies in Oceanography 53: 370-386. 

U.S. Fish and Wildlife Service. 2005. Seabird Conservation Plan Pacific Region. 

U.S. Fish and Wildife Service. 2008. Biological program standard operating procedures (SOPs): French Frigate Shoals, 
Hawaiian Islands National Wildlife Refuge. U.S. Fish and Wildlife Service, Honolulu, Hawaii, USA. 

Weimerskirch, H., Le Corre, M., Jaquemet, S., Potier, M., and R. Marsac. 2004. Foraging strategy of a top predator in 
tropical waters: great frigatebirds in the Mozambique Channel. Marine Ecology Progress Series 275: 297-308. 

Whittow, G.C. 1993. Laysan Albatross (Phoebastha immutabilis), The Birds of North America Online (A. Poole, Ed.). 
Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/ 
species/066 

Whittow, G.C. 1997. Wedge-tailed Shearwater {Puffinus pacificus), The Birds of North America Online (A. Poole, Ed.). 
Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/ 
species/305 

PERSONAL CONTACTS 

Anders, A. Clancy Environmental Consultants. Honolulu, HI, USA. 

Flint, B. U.S. Fish and Wildlife Service, Honolulu, HI, USA. 
Klavitter, J. USFWS, Honolulu, HI, USA. 

WEBSITES 

Hawaii Division of Forestry and Wildlife. 2008. http://www.state.hi.us/dlnr/dofaw/cwcs/Conservation_need.htm#Species 

Pacific Seabird Group and USGS, Alaska Biological Science Center, 2007 Pacific Seabird Monitroing Database, http:// 
www.absc.usgs.gov/research/psinfonet/psmdb/splashpsmdb.htm 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Nonindigenous and Invasive Species 

Kevin See 1 , Scott Godwin 2 and Charles Menza 3 



INTRODUCTION 

The Northwestern Hawaiian Islands (NWHI) represents a relatively pristine marine ecosystem with few non- 
indigenous and invasive species. Of the 343 nonindigenous species (NIS) found in the water's of the Main 
Hawaiian Islands (MHI), only 13 have been detected in the NWHI (Eldredge and Carlton, 2002; Godwin et 
al., 2006; Godwin et al., 2008). This difference is likely due to the NWHI's extreme remoteness, relatively low 
rates of visitation and concerted management efforts. Still, the threat of nonindigenous species spreading from 
the MHI to the NWHI and becoming invasive is a serious concern. The terms nonindigenous and invasive are 
both used to refer to species that are living outside of their historic native range. The difference is that invasive 
species have been shown to cause environmental or economic harm, while NIS have not. Most NIS currently 
found in the NWHI are in few locations and in low abundances. There is debate as to whether any are invasive, 
but this is an active area of research (Schumacher and Parrish, 2005). 

A total of 13 nonindigenous species have been authoritatively detected in the NWHI (Figure 8.1; Table 8.1). 
These species range from invertebrates to fish, and have a wide variety of life histories, likely modes of intro- 
duction and potential impacts. Some species have been found in only one or two locations (e.g., the red alga 
Hypnea musciformis), whereas others are widely distributed throughout most of the atolls and shoals (e.g., the 
blueline snapper Lutjanus kasmira). The difference in their distributions is related to their movement speeds, 
transport methods, ecological success and probability of detection. 




Figure 8.1. Documented distribution of nonindigenous and invasive species in the NWHI. Map: K. Keller. 



1. University of Washington 

2. University of Hawaii 

3. NOAA/NOS/NCCOS/CCMA Biogeography Branch 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 8.1. Marine nonindigenous and invasive species in the Northwest Hawaiian Islands. The table also includes infor- 
mation on their native range, where they have been seen in the NWHI, present population status and potential impacts. 
Sources: Abbott, pers comm; DeFelice et al., 1998; DeFelice et al., 2002; Godwin et al., 2004; Godwin, 2008; Godwin, 
pers comm; Waddell et al., 2008; Zabin et al., 2004. 



SCIENTIFIC 

NAME 


COMMON 
NAME 


TAXA 


NATIVE 
RANGE 


P STATUS T SIGHTINGS POTENTIAL IMPACT 


Hypnea 
musciformis 


Red alga 


Algae 


Unknown; 
Cosmopolitan 


Unknown; in 
drift and on 
lobster traps 


MMM 


Change community structure and 
diversity of benthic habitat, including 
overgrowing coral. Currently forms 
large blooms, up to 7,465 kg (or 
20,000 lbs), off the coast of Maui. 


Diadumene 
lineata 


Orange- 
striped sea 
anemone 


Anemone 


Japan 


Unknown; on 

derelict net 

only 


PHR 


Fouling organism. Ecological impact is 
unstudied but presumed minimal. 


Pennaria 
disticha 


Christ- 
mas tree 
hydroid 


Hydroid 


Unknown; 
Cosmopolitan 


Established 


MMM, FFS, 

GAR, MAR, 

LAY, LIS, PHR, 

MID, KUR 


Competition for space with other inver- 
tebrates. Also stings humans, causing 
a mild irritation. 














the potential to overgrow coral reefs. 


Schizoporella 
errata 


Branching 
bryozoan 


Bryozoan 


Mediterranean 


Established 


MID 


Fouling organism. Ecological impact 
unstudied, but observations suggest 
some competition for space with other 
fouling invertebrates. 


Balanus 
reticulatus 


Barnacle 


Barnacle 


Atlantic 


Established on 
seawall 


FFS 


Fouling organism. Ecological impact is 
unstudied but presumed minimal. 


Balanus 
venustus 


Barnacle 


Barnacle 


Atlantic and 
Caribbean 


Not estab- 
lished; on ves- 
sel hull only 


MID 


Fouling organism. Ecological impact is 
unstudied but presumed minimal. 


Chthamalus 
proteus 


Caribbean 
barnacle 


Barnacle 


Caribbean 


Established in 
harbor 


MID 


Serious nuisance fouling organism. 
Competes for space and food resourc- 
es with native species. Grows in such 
densities that it could exclude algal 
grazers such as opihi. 
















Polycarpa 
aurita 


Styelidae, 
solitary 
tunicate 


Tunicate 


Indo-Pacific, 
Western 
Atlantic 


ARMS 


FFS 


This species has the capacity to 
become an aggressive component of 
a fouling community on man-made 
surfaces, and the potential for recruit- 
ment to natural habitats is always a 
possibility. 


Lutjanus 
fulvus 


Toau or 
Blackline 
Snapper 


Fish 


Indo-Pacific 


Established 


FFS 


Could out-compete native species for 
resources, but current densities may 
be too low to see these effects. 


Lutjanus 
kasmira 


Taape or 
Blueline 
snapper 


Fish 


Indo-Pacific 


Established 


NIH, FFS, 

MAR, LAY, 

MID 


Could prey on or out-compete desir- 
able fishery species. May also exclude 
more desirable species from fishing 
gear through competition. Scientific 
research into these effects is currently 
lacking. 


Cephalopho- 
lis argus 


Roi or 
Peacock 
grouper 


Fish 


Indo-Pacific 


Established 


NIH, MMM, 
FFS 


May predate on native species that 
are targeted by aquariums, dive tours 
and fishermen. Scientific research into 
these effects is currently lacking. 


Carijoa riisei 


Snowflake 
coral 


Octocoral 


Indo-Pacific 


Has not been 

seen in NWHI 

yet 


Five Fathom 
Pinnacle 


Overgrows black corals, killing them. 
Competes for space with other inver- 
tebrates. 


Acanthophora 
spicifera 


Red alga 


Algae 


Indo-Pacific 


Has not been 

seen in NWHI 

yet 


Kauai 


Change community structure and 
diversity of benthic habitat, including 
overgrowing coral. 


Island/atoll abbreviations found throughout this chapter: NIH = Nihoa, MMM = Mokumanamana, FFS = French Frigate Shoals, GAR = Gardner 
Pinnacles, MAR = Maro Reef, LAY = Laysan Island, LIS = Lisianski Island, PHR = Pearl and Hermes Atoll, MID = Midway Atoll, KUR = Kure Atoll 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

All of the atolls and islands have at least one nonindigenous species, but several such as Midway Atoll (six 
species) and French Frigate Shoals (five species) have numerous. These two locations have been the foci of 
human activity for many years, especially during World War II when they were used as military bases. This ac- 
tivity probably meant greater ship traffic and food imports, both of which are considered principal NIS vectors. 
They are also two of the most studied locations and thus present NIS have a greater probability of detection. 

In addition to confirmed NIS observations in the NWHI, several unconfirmed reports of sightings exist and 
two other species (i.e., Carijoa riseii and Acanthophora Spicifera) have proven to be extremely successful 
invaders of the MHI, and therefore pose a serious threat to the NWHI. The red algae Hypnea musciformis and 
Acanthophora spicifera may have been sighted drifting on Maro Reef and sighted near Midway Atoll, respec- 
tively. The blackline snapper (Lutjanus fulvus) may have been spotted off Nihoa Island, and blueline snapper 
(L. kasmira) may have been seen off Mokumanamana, Lisianski Island, and Pearl and Hermes Atoll (Godwin 
et al., 2006, Draft Environmental Impact Statement, Draft Management Plan for the NWHI Proposed National 
Marine Sanctuary 2006, R. Kosaki, pers. comm.). 



Vectors 

Populations of nonindigenous marine species that have already colonized areas of the MHI represent the most 
likely source of nonindigenous species in the NWHI. This deduction is based on the proximity and pattern of 
ship movements among these two areas (Godwin et al., 2006). It is difficult to conclusively determine vectors 
of movement, but the most likely are: hull fouling, ballast water discharge and natural water currents. Recently, 
marine debris has been suggested as a vector and has shown the ability to transport nonindigenous species 
to the NWHI (Godwin et al., 2006). To date no records show any species were purposefully introduced into the 
NWHI, although they most certainly were to the MHI (e.g., blueline and blackline snapper, Peacock grouper). 

Data Collection 

To deal with the threat of NIS and invasive species, information about their biology and spatial distribution is 
critical. Sightings of marine invasive species in the NWHI come from a variety of sources (Table 8.2). Sources 
are typically biological inventories of particular areas (e.g., Midway Harbor Survey, French Frigate Shoals Sur- 
vey) or are opportunistic (e.g., derelict fishing net removal project) and thus are limited in temporal and spatial 
scope. These types of data are useful for determining if a particular location has been invaded, or if a potential 
vector is acting as an invasive pathway. However, these data do not provide any indication of the severity of an 
invasion, whether an invasive population is growing or shrinking or the ability to complete a rigorous statistical 
comparison among locations. 

Currently, there is no systematic survey which covers all habitats likely to harbor NIS and invasive species. 
Most data are collected or informed by conventional SCUBA or snorkeling. As a result, most data are collected 
at depths shallower than 35 m. This is a concern since several nonindigenous species already detected in the 
NWHI or in the MHI have been detected well below this limit (e.g., blueline snapper - 256 m). To fill this gap 
Papahanaumokuakea Marine National Monument (PMNM) has begun assessing deep water survey technolo- 
gies (C. Menza, pers. comm.). 

The NWHI Coral Reef Assessment and Monitoring Program (NOWRAMP) and lobster trap monitoring pro- 
grams provide quantitative abundance data of NIS and can monitor changes over time (see Table 8.2 for 
details); however sampling is spatially biased. For example, NOWRAMP surveys are completed at permanent 
sites and thus may not be representative of larger populations and may not detect NIS that occur in unsampled 
habitats. Similarly, hull, net and trap inspections are tied to the distribution of invasive species and may provide 
biased population estimates of attached species. More intensive surveys in specific areas (e.g., Midway Har- 
bor Survey) offer detailed fine spatial scale data and taxonomic resolution, but are time intensive and costly. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 8.2. Marine invasive species monitoring programs in the NWHI. 



PROGRAM 


OBJECTIVES 


TIME 
PERIOD 


ISLANDS OR 
ATOLLS 


AGENCIES 


NOWRAMP 


Monitor fish, algae, coral and other 
invertebrates 


2000-2007 


NIH, MMM, 

FFS, GAR, 

MAR, LAY, LIS, 

PHR, MID, KUR 


NOAA-PMNM, 
PIFSC 


Midway Harbor Survey 


Survey the invertebrates on artificial 
substrates in and around Midway Harbor 


1998 


MID 


USFWS, 

Bishop 

Museum 


French Frigate Shoals 
Survey 


Survey the seawall at Tern Island for 
nonindigenous species 


2002 


FFS 


USFWS, 

Bishop 

Museum 


Derelict Fishing Net 
Removal Project 


Remove derelict fishing nets on Kure, Pearl and 
Hermes, Midway and Lisianski and determine if 
any nets contained nonindigenous species 


2000 


LIS, PHR, MID, 
KUR 


NOAA-NMFS 


Derelict Fishing Net 
Removal Project 


Remove derelict fishing nets on French Frigate 
Shoals and determine if any nets 
contained nonindigenous species 


2007 


FFS 


NOAA-NMFS 


Census of Coral Reefs 


Characterize invertebrate communities 


2007 


FFS 


NOAA-NMFS 


Hull Fouling Project 


Assess hull fouling as a mechanism for the disper- 
sal of nonindigenous species 


2003 


MHI, MID 


HCRI-RP, 
HI-DLNR 


Lobster Trap Monitoring 


Monitor the population of spiny lobsters, and iden- 
tify any algae that is growing on the 
lobster traps 


1985-2007 


MMM, MAR 


NOAA-PIFSC 


Abbreviations: NOWRAMP = Northwest Hawaiian Islands Rapid Assessment and Monitoring Program, MHI = Main Hawaiian Islands, NOAA = 
National Oceanic and Atmospheric Administration, PMNM = Papahanaumokuakea Marine National Monument, USFWS = U.S. Fish and Wildlife 
Service, NMFS = National Marine Fisheries Service, HI DLNR = Hawaii Department of Land and Natural Resources, PIFSC = Pacific Islands Fish- 
eries Science Center, HCRI-RP = Hawaii Coral Reef Initiative Research Program 



MARINE ALGAE 

Nonindigenous algae in the NWHI are a major concern, because of the mobility of propagules, fast growth 
rate, potential ecological impacts to the native benthic community and presence in the MHI. One species of 
red algae, Hypnea musciformis, has been detected in the NWHI and another species, Acanthophora spicifera, 
is of particular concern because of its aggressive growth rate. Both species are present in the MHI and H. 
musciformis probably originated there. 

At least 19 species of macroalgae have been intentionally or passively introduced in Hawaii since the mid 
1950s (Doty, 1961; Brostoff, 1989; Rodgers and Cox, 1999; Russell, 1987, 1992; Woo, 1999; Smith et al., 
2002; Smith et al., in press) and at least five have successfully established themselves. These species are 
capable of moving to the NWHI. 



Red Algae, Spiny Algae (Acanthophora spicifera) 

This species of red algae has not yet been authoritatively recorded in the NWHI, but there has been one un- 
confirmed sighting at Midway and due to its success in the MHI, it is a species of particular concern. It is widely 
distributed among the MHI and throughout the tropics and subtropics. Introduction likely originated in Honolulu 
Harbor in the 1950s via a fouled barge originating in Guam (Doty, 1961). It has since spread to all the MHI, and 
is the most widespread invasive algae in the archipelago and is now a common component of the intertidal 
community (Smith et al., 2002). 

Movement and associated range extensions occur naturally through water movement, or anthropogenically 
through hull fouling. Fragments or spores move through advection and are likely the means of local dispersal 
in Hawaii (Kilar and McLachlan, 1986). Branches are brittle that often results in fragmentation. Fragments 
can accumulate forming large, free-floating populations and can drift for potentially long distances before set- 
tling and establishing new colonies. It is also frequently spotted fouling hulls throughout the MHI (Smith et al., 
2002). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

A. spicifera can adapt to a variety of habitats and environmental conditions, and this is one of the reasons of its 
success throughout tropical and subtropical ecosystems. In Hawaii, it is abundant in protected areas where it 
is not exposed to high-energy wave action, such as rocky intertidal beaches, tide-pools and shallow reef-flats. 
It attaches to hard substrates and is often found growing with the native algae species of Laurencia nidifica 
and Hypnea cervicornis (Botany UH, 2001). In other areas it has been found as an epiphyte on other algae 
species and as a free living drift alga. 

Potential impacts are poorly studied. It likely impacts the community structure and diversity of the benthic 
habitat through competition and smothering (Preskitt, 2002; Eldrege 2003), but these effects have not been 
well quantified (Shluker, 2003). A. spicifera can outcompete native algae such as L. nidifica and H. cervicornis 
(Russell, 1992). In the eastern tropical Pacific, blooms of A. spicifera covered by cyanobacterial epiphytes 
have been observed at several reefs and were associated with widespread coral mortality. 



Red Algae (Hypnea musciformis) 
In 2005, international press coverage 
drew attention to the potential spread of 
the red, invasive alga, Hypnea musci- 
formis when large quantities were found 
entangled in lobster traps at depths from 
30 to 90 m near Mokumanamana (God- 
win et al., 2006; Figure 8.2). The spe- 
cies was first recorded from deep water 
(>30 m) at Mokumanamana in 2002, 
and one small individual was found as 
part of a drift assemblage at Maro Reef 
(Friedlander et al., 2008). From 2002 
through 2004, small sprigs of the alga 
were commonly recorded on lobster 
traps at Mokumanamana. In spring to 
early summer of 2005, pounds of H. 
musciformis began to appear on lobster 
traps at Mokumanamana, generating 
concern about a large-scale epidemic 
of this nuisance alga. Later that year a 
special cruise was organized by PMNM 
to investigate the problem. Interestingly, 
no H. musciformis was discovered at 
Mokumanamana during the cruise, and 

continued investigations of algae associated with lobster traps in 2006 have failed to find any significant popu- 
lation blooms other than a few small individuals similar to those documented in 2002 through 2004 (Fried- 
lander etal., 2008). 




Figure 8.2. General location of the red algae Hypnea musciformis from 
NOAA/PIFSC lobster trap monitoring. 



H. musciformis was intentionally introduced from its native range in Florida to Kaneohe Bay on Oahu in 1974 
for mariculture. It is commercially cultivated as a food source and for kappa carrageenan, a common food ad- 
ditive. Like A. spicifera, it spreads quickly and is distributed widely throughout the MHI where it is now found 
on Kauai, Oahu, Molokai and Maui, with the most abundant populations occurring on Maui (Botany UH, 2001). 
Populations are often found on calm intertidal and shallow subtidal reef-flats where it either attaches to sandy 
flat rocks or is found as an epiphyte on other algae species, often on A. spicifera, Laurencia nidifica, Sargas- 
sum echinocarpum, and S. polyphyllum (http://hawaii.edu/reefalgae/invasive_algae/index.htm). 



Principal reasons for this species success are its high growth rate, ability to epiphytize other algae and fre- 
quent fragmentation. Russell (1992) estimated a growth rate between 10-50% per day. Drifting fragments 
can attach to other floating algae, like S. echinocarpum or S. polyphyllum, and float long distances before es- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

tablishing new colonies. Attachment is 
aided by the presence of apical hooks 
(Figure 8.3). Fragments as small as 5 
mm proved viable, growing at a rate of 
200% a week (Smith et al., 2002). Be- 
sides fragmentation, H. musciformis 
also spreads through hull fouling. 

Potential impacts include competition 

with native algae and the creation of 

large dense surface mats. Like other 

invasive algae, it probably impacts the 

community structure and diversity of the 

benthic habitat, but these effects have 

not yet been quantified (Shluker, 2003). 

Russell (1992) found H. musciformis 

can outcompete the native algae H. cer- 

vicornis, especially in the presence of A. spicifera. H. musciformis can form large dense mats, which have 

been correlated with high levels of nutrient inputs from the coast. Similar nutrient inputs are not present in the 

NWHI, but mats located around the MHI are capable of supplying propagules for distribution to the NWHI. The 

presence of dense mats are also a concern, because in peak blooms tens of thousands of pounds of algae 

can wash ashore forming windrows 0.5 m high. The effect of these windrows on local biota like the Hawaiian 

monk seal or green sea turtle is unknown. 

H. musciformis now makes up a significant portion of the diet for the green sea turtle, sometimes composing 
as much as 99-100% of the seaweed mass in their stomachs. However, the nutritional value of H. musciformis 
has not yet been determined and so the long-term impact of incorporating this alga into the sea turtles' diet is 
unknown (Botany UH, 2001). 















'\& 




^^^ JWi 






■'i K vL 



Figure 8.3. H. musciformis. 
hooks. Photo: P. Vroom. 



The arrows point to the species' distinctive 



INVERTEBRATES 

Out of the all the different taxonomic groups of NIS, invertebrates represent the most species and are the least 
studied. Nine invertebrate species (one anemone, one hydroid, two bryozoans, three barnacles and two tuni- 
cates) have been detected in the NWHI. These invertebrates are typically cryptic and have been detected with 
the help of fine-scale surveys in targeted areas (e.g., Defelice et al., 1998, 2002). Most nonindigenous inverte- 
brates have been detected at Midway Atoll and French Frigate Shoals, the two locations with the lion's shares 
of survey effort and human activity. A tenth invertebrate species, the snowflake coral (Carijoa riseii), which has 
not been detected in the NWHI is described herein because it is a species of particular concern. 



Orange-striped Sea Anemone (Diadumene lineata) 

The orange-striped sea anemone is native to Japan, but has spread throughout the Pacific, Atlantic, Carib- 
bean, the North Sea and the Mediterranean (Zabin et al., 2004). In 2000, about 100 individuals were identified 
in the lagoon at Pearl and Hermes Atoll attached to a derelict fishing net (Zabin et al., 2004; Figure 8.4). To 
date, no established adults have been seen in the NWHI. 

Although it can reproduce sexually, it likely spreads through asexual reproduction and hull fouling in the NWHI 
(Zabin et al., 2004). It exhibits a wide tolerance of temperature and salinity and is generally found on solid 
substrates, in intertidal pools or protected shallow waters such as bays and harbors. The orange-striped sea 
anemone is often found with mussels and oysters in other parts of its range (DeFelice et al., 2001), and could 
have been transported to Hawaii in an oyster shipment (Zabin et al., 2004). The impacts of this species in the 
NWHI remain unknown and unstudied. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Christmas Tree Hydroid (Pennaria 
disticha) 

The Chrismas tree hydroid is native to 
the western Atlantic and has been re- 
ported in all of the NWHI except Nihoa 
(Godwin et al., 2006). It also is widely 
distributed among the MHI (DeFelice et 
al., 2001). It was first reported in the re- 
gion during a survey of Pearl Harbor in 
1929 (DeFelice et al., 2001). 

It attaches to natural and artificial hard 
substrates where there is some water 
movement. It is very common in har- 
bors in all the MHI and is often found 
in more protected areas such as cracks 
and crevices on reefs, at depths of 0-50 
m. The impacts of the Christmas tree 
hydroid are unstudied, but it is likely that 
it competes for space with other inver- 
tebrates. It also can sting humans, re- 
sulting in minor irritation (DeFelice et al., 
2001). 




Figure 8.4. General location of the orange-striped sea anemone (Diad- 
umene lineata) from NOAA/PIFSC/CRED Marine Debris Program. 



Bushy Bryozoan (Amathia distans) 
In 1997 the bushy bryozoan was found 
at Midway Harbor, dominating many 
of the manmade structures that were 
surveyed (Figure 8.5). It formed large 
colonies on wood, concrete and metal 
pilings, as it does in harbors in the MHI 
(DeFelice et al., 1998). To date, this is 
the only location in the NWHI where it 
has been sighted. Its native range is the 
Caribbean, but it has spread over much 
of the tropics and subtropics including 
the western Atlantic, Mediterranean and 
Red Seas, eastern Pacific and coastal 
waters of Australia, New Zealand, Java 
and Japan (DeFelice etal., 2001). Move- 
ment is considered to be aided by hull 
fouling, ballast water discharge (larvae) 
or natural water movement (Shluker, 
2003). 




Figure 8.5. General locations of the bushy bryozoan (Amathia distans) at 
Midway Atoll. 



The bushy bryozoan was first spotted in the region at Kaneohe Bay in 1935, and has since spread to all the 
MHI (Shluker, 2003; Coles et al., 2004). It can be found in shallow water on hard anthropogenic substrates 
such as pilings and vessel hulls and natural substrates such as coral rubble. It is usually found inside harbors 
or embayments, or occasionally in more protected areas of the reef. The impacts of the bushy bryozoan are 
unknown and presumed minimal (DeFelice et al., 2001), probably including competition for space (Shluker, 
2003). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Branching Bryozoan (Schizoporella errata) 
The branching bryozoan was recorded 
at Midway Harbor in 1997, where it was 
found occupying many of the same lo- 
cations as the bushy brozoan, although 
not as abundant (DeFelice et al., 1998) 
(Figure 8.6). It is usually found inside 
harbors or embayments on man-made 
substrates, or occasionally in more pro- 
tected areas of coral reefs (DeFelice et 
al., 2001). Its native range is the Medi- 
terranean, but is now found worldwide, 
including all the MHI (DeFelice et al., 
2001) where it was first described at Pearl 
Harbor in 1933. It can be transported 
anthropogenically through hull fouling, 
which is likely how it was unintentionally 
transported to so many locations around 
the globe (Shluker, 2003). The impacts 
of this species are unknown, but likely 
include competition for space (DeFelice 
etal.,2001). 




Legend 
Invertebrates 

Branching bryozoan 



Figure 8.6. General locations of the branching bryozoan (Schizoporella er- 
rata) at Midway Atoll. 



Barnacle (Balanus reticulates) 
Although this species of barnacle has 
been found in the MHI on Kauai, Oahu, 
Maui and Hawaii (Coles et al., 2004), 
and was found on about 25% of the ship 
hulls in one hull fouling study (Godwin 
et al., 2004), it has only been spotted 
once in the NWHI, on a seawall at Tern 
Island in French Frigate Shoals in 2002 
(DeFelice et al., 2002; Figure 8.7). It is a 
fouling organism. Its ecological impact is 
presumed to be minimal, although there 
is little research to confirm this assump- 
tion. 




Balanus reticulatus at Tern 



Barnacle (Balanus venustus) 
This barnacle, native to the Atlantic and 
Caribbean oceans, has been seen once 
on a hull of a ship anchored at Midway 
Harbor in 2003 (Godwin et al., 2004), 
demonstrating this species' ability to be 

transported through hull fouling. However, an established adult has never been seen in the NWHI. Its ecologi- 
cal impact is presumed to be minimal. 



Figure 8. 7. General location of the barnacle 
Island, French Frigate Shoals. 



Caribbean Barnacle (Chthamalus proteus) 

This barnacle from the Caribbean was found in Midway Harbor attached to pier pilings in 1997 (DeFelice et al., 
1998; Figure 8.8). It likely arrived in the region between 1973 and 1994, since it was first noticed at Kaneohe 
Bay, Oahu in 1995 and was not found during a comprehensive intertidal survey of Oahu in 1972. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Figure 8.8. General locations of the Caribbean barnacle fChthamalus pro- 
teus) at Midway Atoll. 



It was probably introduced through ei- 
ther hull fouling or ballast water, although 
Southward et al. (1998) argues that hull 
fouling is more likely. It is commonly seen 
above the waterline on inter-island ships 
(Zabin, 2007). Larval dispersal could 
also be a natural vector for spread be- 
tween the islands of the Hawaiian archi- 
pelago, now that it is established there. 
Although there may be some peaks of 
larval production, larvae are found in the 
water column year-round. 

Surveys of MHI have found Caribbe- 
an barnacles around Kauai, Maui and 
Hawaii (DeFelice et al., 2001). It usu- 
ally colonizes supratidal anthropogenic 
structures such as pier pilings and sea 
walls, although some individuals have 
been observed on intertidal boulders in 
the MHI. It is generally found in protect- 
ed embayments and harbors, but small colonies have been found at one high energy site in Kaneohe Bay. This 
finding is a concern, because this species may be moving into habitat used by the native barnacle Nesoch- 
thamalus intertextus. At the moment, it seems the Caribbean barnacle is not competing with N. intertextus, 
but rather growing next to it. In addition, Caribbean barnacle individuals were quite small, so it was unclear 
whether there was an established population. 

The Caribbean barnacle has been implicated in displacing another nonindigenous barnacle, Balanus amphi- 
trite, in the MHI demonstrating its competitive ability (Shluker, 2003). Its rapid proliferation may reflect that it is 
filling an unexploited niche in the Hawaiian archipelago, in the high intertidal and splash zones. The density of 
colonies and the rapid pace of reproduction make the Caribbean barnacle a good competitor for space. This 
proliferation could alter the community structure and potentially exclude algal grazers such as protected Ha- 
waiian limpets (e.g., Cellana exarata, C. melanostoma, C. sandwicensis, C. talcosa). 

Styelidae, Solitary Tunicate (Cnemidocarpa Irene) 

This species is a widespread Indo-Pacific tunicate found in Japan, the Philippines, Australia, Micronesia and 
Melanesia. Large specimens may reach a length of 4 cm and have a dark brown to whitish tunic with deep 
wrinkles that are arranged to create irregularly shaped raised areas. This species is commonly associated with 
fouling communities located within man-made harbors and shallow benthic habitats with rubble substrate from 
Kauai to the island of Hawaii (Abbott et al., 1997). 

The larval stage of most solitary tunicates is brief; the larva does not feed, but concentrates on finding an ap- 
propriate place for the adult to live. The actual larvae are tadpole shaped and the muscular tail comprises two- 
thirds of the larval body; it is supported by a notochord and contains a nerve cord. Gravity and light-sensitive 
sensory vesicles along the dorsal surface of the larval body orient the animal as it swims. After a period of up 
to a few days, the larva will settle and attach itself to a surface using three anterior adhesive papillae. As the 
larva metamorphoses into an adult, the tail reabsorbs, providing food reserves for the developing animal. 



This species has only been recorded from French Frigate Shoals in the Monument, where it was collected from 
an Autonomous Reef Monitoring Structures (ARMS) installed in 2006 (Godwin etal., 2008; Figure 8.9). Due to 
the short larval duration of tunicates, this species was likely transported to French Frigate Shoals by some an- 
thropogenic means from a source location in the southeastern portion of the archipelago. Therefore this record 
represents recruitment to the ARMS from an undocumented established population at French Frigate Shoals. 





Figure 8.9. Documented location ofC. irene at French Frigate Shoals. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

The impacts of this species are unknown \%jz 
but it has the capacity to become a domi- 
nant fouling organism on any man-made 
substrate. 



Styelidae, Solitary Tunicate (Polycar- 
pa aurita) 

This solitary tunicate is pale brown with 
a tough and leathery tunic that is gener- 
ally encrusted with worm tubes, sponges 
and other fouling organisms. Specimens 
in Hawaii only reach up to 4 cm in length 
but this species attains greater lengths 
(10-12 cm) in other areas of its Indo-Pa- 
cific range. This species is also found in 
the western Atlantic (Caribbean and Gulf 
of Mexico). It is established in the south- 
eastern portion of the archipelago as a 
common species in fouling communities 

located within man-made harbors and the shallow and intertidal habitats of natural embayments (Abbott et al. 
1997). 

The larval cycle described under C. 
irene also applies to this species. There- 
fore, a larval cycle of only a few days 
exists. It was recently recorded from 
French Frigate Shoals from the same 
collections in which C. irene was identi- 
fied (Godwin et al., 2008). These collec- 
tions were part of an effort by the Coral 
Reef Ecosystem Division (CRED) of the 
Pacific Islands Fisheries Science Cen- 
ter in Honolulu in 2007. The focus of the 
efforts was to expand a 2000 project, 
which examined fouling organisms as- 
sociated with derelict fishing gear in the 
NWHI (Godwin, 2000; Figure 8.10) and 
retrieve and quantify the organisms col- 
lected by an ARMS deployed in 2006 at 
French Frigate Shoals. As with C. irene, 
anthropogenic transport to French Frig- 
ate Shoals is assumed and a scenario of opportunistic recruitment to the ARMS from some established popu- 
lation in the lagoon is likely. 

This species has the capacity to become an aggressive component of a fouling community on man-made sur- 
faces, and the potential for recruitment to natural habitats is always a possibility. Recent incidences of natural 
tunicate populations acting invasively and overgrowing remote coral reef areas demonstrates the potential of 
this group of organisms to cause damage to coral reefs without direct human influence (Littler and Littler, 1995; 
Vargas-Angel et al., 2008) 




Figure 8.10. Documented location of P. aurita at French Frigate Shoals. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Snowflake Coral (Carijoa riisei) 

The snowflake coral has not been detected in the NWHI, but is a species of particular concern. It was first spot- 
ted in Pearl Harbor in 1972 (DeFelice et al., 2001), and by 1990 had been recorded around all of the MHI. Of 
the 343 nonindigenous marine species that have been introduced to the Hawaiian Islands, the snowflake coral 
may be the most successful at proliferation, as demonstrated by its distribution among the MHI, and it may 
exhibit some of the highest invasive potential (Grigg, 2003). It has not been sighted in the NWHI to date, but 
in 2007 a colony was found at Five Fathom Pinnacle (Kahng, per comm.), approximately 200 km from Nihoa 
Island which is the southeastern-most point of the NWHI. 

This species was originally thought to be native to the Caribbean, but recent research has shown it to be more 
likely indigenous to the Indo-Pacific. It is likely that several slightly different species have reached the Hawai- 
ian archipelago (Kahng, 2006). 

The snowflake coral is very light sensitive; it thrives in spots that receive 10-30% ambient light, and avoids 
well-lit habitats. Therefore in shallow water (10-30 m), where light levels are high, it attaches to dark cracks, 
shaded walls or pilings, the underside of ledges and corals, lava tubes and other shaded areas. As it moves 
into deeper water and light levels diminish, it is found on a wider variety of habitats. At depths of 75-110 m, 
it has been found to explode into patches as large as 200 km 2 (Grigg, 2003). It generally attaches to hard 
substrates such as rocks, corals or anthropogenic structures. It does need to be positioned above the benthic 
layer, and away from stagnant water, as it requires some wave energy to continuously transport the zooplank- 
ton that it filters from the water for food (Godwin et al., 2006). 

The snowflake coral reproduces both asexually and sexually. The polyps can split in two, allowing clones to 
spread and cover an entire habitable patch within several years. It can also release gametes into the water 
column, which once fertilized, can survive for up to 90 days (Kahng, 2006) and thus are capable of travelling 
long distances. This species can also spread through hull fouling, although this may not be common. 

At shallow depths, the snowflake coral seems to occupy an unutilized habitat niche in Hawaii (Shluker, 2003). 
However at depth, it has overgrown entire beds of black coral, killing 90% of the coral surveyed in the Maui 
Black Coral Bed in 2001 (Grigg, 2003). Black coral harvesting generates $15 million a year in the state of Ha- 
waiian, and the spread of the snowflake coral represents a serious threat to this industry (Godwin et al., 2006). 
Beyond the economic impacts, it has shown the potential to severely reduce biodiversity by blanketing entire 
areas. 



FISHES 

Three species of nonindigenous fish have been observed in the NWHI, blackline snapper (Lutjanus fulvus), 
blueline snapper (L. kasmira) and Peacock grouper (Cephalopholis argus). All three species were purposefully 
introduced to the MHI between 1955 and 1961 along with eight other species of groupers (Serranidae), snap- 
pers (Lujanidae) and emperor breams (Lethrinidae) from Moorea in French Polynesia. All were introduced as 
potential commercial species (Brock, 1960; Randall, 1987). Of the three species, blueline snapper have been 
the most successful in terms of distribution and abundance (Shluker, 2003). 



Blackline Snapper (Lutjanus fulvus or Toau) 

Intentionally introduced in 1956, blackline snapper has spread to all of the MHI, and into the southeastern 

end of the NWHI. It has been spotted at Nihoa and French Frigate Shoals (Shluker, 2003; Figure 8.11). It has 

fairly low abundance, possibly due to its exploitation for food (Shluker, 2003). Blueline snapper (L. kasmira) 

was introduced around the same time, but it has spread much faster than blackline snapper, despite the many 

biological similarities between the two species. Scientists are unsure how to explain the difference in range 

expansion. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Blackline snapper is a reef fish, gener- 
ally found in the lagoons or outer reef 
slopes and usually at depths of 1-40 m, 
but it has been seen as deep as 75 m. It 
has a temperature tolerance of 20-28°C 
and spawns year-round (http://www.lar- 
valbase.org), increasing its chances of 
larval dispersal. Ecological impacts are 
unstudied. 




manamana 



French 
Frig 



Ihoals 



Nihoa 
'and 



Legend 

Fish 




Figure 8.11. Documented distribution of blackline snapper (Lutjanus 
fulvusj in the NWHI. 



Blueline Snapper 
(Lutjanus kasmira or Taape) 
Blueline snapper has been detected 
throughout the NWHI, including Nihoa, 
Mokumanamana, French Frigate Shoals, 
Maro Reef, Laysan Island and Midway 
Atoll (Friedlander et al., 2005). It likely 
migrated from the MHI where it was in- 
tentionally introduced to Oahu in 1955. 
From the initial population of 3,200 indi- 
viduals brought from French Polynesia, 
the fish has spread throughout the full 
length of the Hawaiian archipelago (Oda 
and Parrish, 1982; Randall et al., 1993; 
Figure 8.12) and is now one of the most 
conspicuous and abundant species in 
the fish community. Friedlander et al. 
(2002) found blueline snapper was the 
second most abundant species by num- 
ber and biomass over hard substrate in 
Hanalei Bay, Kauai. 

Due to its abundance and the concern 
that blueline snapper might impact na- 
tive fish, more effort has been spent 
studying its ecology compared to other 
similar nonindigenous species. Blueline 
snapper is generally found in lagoons 

and outer reef slopes at depths from 2-70 m, but it has been seen as deep as 256 m. Friedlander et al. (2002) 
found the species to be abundant over habitats like deep slope, spur and groove and shallow slope, but it was 
also found in lesser quantities in the complex back reef. A more recent report indicated that blueline snapper 
is also common among algal plain habitats (C. Menza, pers. comm.). These low relief habitats dominated by 
algae (macroalgae and crustose coralline algae), may make up a considerable proportion of the deeper ben- 
thic habitats in the NWHI where coral are rare. Friedlander et al. (2002) have also shown that blueline snapper 
utilize sand habitats for feeding and the species may undergo an ontogenetic habitat shift. 

The blueline snapper was never accepted into the local diet, and many fishermen believe it out competes na- 
tive fish for resources and fishing bait. There is little scientific evidence to back this conclusion (but see Schu- 
macher and Parrish, 2005), which leads to disagreement and debate between scientists and fishermen as to 
the effects of the blueline snapper on native species (Shluker, 2003). 




Figure 8.12. Documented distribution of blueline snapper (Lutjanus kas- 
mira) in the NWHI. Source: Sladek Nowlis and Friedlander, 2004. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Peacock Grouper (Cephalopholis argus or Roi) 

The Peacock grouper was introduced 

from French Polynesia in 1956 as a 

food species. Since then, it has spread 

throughout the MHI, and has been seen 

at Nihoa, Mokumanamana and French 

Frigate Shoals in the NWHI (Shluk- 

er, 2003; Godwin et al., 2006; Figure 

8.13). 

It is found in lagoons and seaward reef 
habitats, at depths of 1-40 m, although 
it generally prefers depths of 10 m or 
less (Godwin et al., 2006). 



Although originally sought by fishermen, 
its popularity declined after incidences 
of ciguatera poisoning increased and 
is now considered by many fishermen 
as unsafe to eat (Godwin et al., 2006). 
Without fishing pressure, the Peacock 
grouper has grown abundant and could 
impact native reef fishes through preda- 




Figure 8.13. Documented distribution of the Peacock grouper (Cephalop- 
holis argus) in the NWHI. Source: Sladek Nowlis and Friedlander, 2004. 



tion as well as competition for space and resources. However, there is little scientific research on the effects 
due to Peacock grouper, and thus no conclusive evidence has been gathered. 



MANAGEMENT 

PMNM has taken active steps to mitigate the threats of NIS, including ballast discharge prohibition, hull in- 
spections and cleaning, snorkel/dive gear treatment and luggage inspection of air passengers. Action plans 
consisting of multiple strategies and activities address PMNM priority management needs. One of the PMNM's 
22 action plans is "to detect, control, eradicate where possible, and prevent the introduction of alien species 
into the Monument". PMNM has also undertaken research to develop knowledge of baseline conditions and 
detect NIS introductions. Early detection greatly increases the probability of NIS control and possibly eradica- 
tion (e.g., Pyne, 1999). 



EXISTING DATA GAPS 

The primary data gap for nonindigenous and invasive species in the NWHI is a complete survey of nonindig- 
enous species across habitats. Surveys need to have a greater spatial distribution to have a more complete 
picture of the nonindigenous and invasive species populations. The following are key datasets needed for 
management and future research efforts: 

• Species inventory; 

• Population size; 

• Rate of spread; 

• Spatial distribution; and 

• Habitat requirements and natural history information for established populations to use in habitat suit- 
ability models. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

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waii. Bishop Museum Special Publication. 64 (6B). 64 pp. 

Botany, University of Hawaii Manoa. 2001. Algae: Invasive alien Acanthophora spicifera. Available from Internet URL: 
http://www.hawaii.edu/reefalgae/invasive_algae/rhodo/acanthophora_spicifera.htm. 

Botany, University of Hawaii Manoa. 2001. Algae: Invasive alien Hypnea musciformis. Available from Internet URL: http:// 
www.hawaii.edu/reefalgae/invasive_algae/rhodo/hypnea_musciformis.htm. 

Brock, V. 1960. The introduction of aquatic animals into Hawaiian water. Int. Revue Hydrobiol. 45: 463-480. 

Brostoff, W. 1989. Avrainvillea amadelpha (Codiales, Chlorophyta) from Oahu Hawaii. Pac Sci 43: 166-9. 

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DeFelice, R.C., S.L. Coles, D. Muir, and L.G. Eldredge. 1998. Investigation of the marine communities of Midway Harbor 
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DeFelice, R.C., L.G. Eldredge, and J.T Carlton. 2001. Nonindigenous marine invertebrates. Contribution No. 2001-005 to 
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DeFelice, R.C., D. Minton, and L.S. Godwin. 2002. Records of shallow-water marine invertebrates from French Frigate 
Shoals, Northwestern Hawaiian Islands, with a note on non-indigenous species. Report to the U.S. Fish and Wildlife Ser- 
vice. Bishop Museum, Hawaii Biological Survey, Bishop Museum Technical Report No. 23. 

Doty, M. S. 1961. Acanthophora, a possible invader of the marine flora of Hawaii. Pac. Sci. 15(4): 547- 552. 

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Eldredge, L.G. 2003. Coral Reef Invasions. In: De Poorter, M. (Ed.). 2003. Aliens (17): 9. 

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211-212. 

Eldredge, L.G. 2005. Assessment of the Potential Threat of the Introduction of Marine Nonindigenous Species in the 
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tion No. 2005-001 to the Hawaii Biological Survey. 

Friedlander, A.M, G. Aeby, S. Balwani, B. Bowen, R. Brainard, A. Clark, J. Kenyon, J. Maragos, C. Meyer, P. Vroom, and J. 
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and A.M. Clarke (eds.), The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 
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Biogeography Team. Silver Spring, MD. 569 pp. 

Friedlander, A.M, G. Aeby, R. Brainard, A. Clark, E. DeMartini, S. Godwin, J. Kenyon, R. Kosaki, J. Maragos, and P. 
Vroom. 2005. The State of the Coral Reef Ecosystems of the Northwestern Hawaiian Islands. Pp. 270-311. In: J. Wad- 
dell (ed.), The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2005. NOAA 
Technical Memorandum NOS NCCOS 11. NOAA/NCCOS Center for Coastal Monitoring and Assessment's Biogeography 
Team. Silver Spring, MD. 522 pp. 

Friedlander, A.M., J.D. Parrish and R.C. Defelice. 2002. Ecology of the introduced snapper Lutjanus kasmira (Forsskal) 
in the reef fish assemblage of a Hawaiian bay, Journal of Fish Biology 60(1): 28-48. 

Godwin, L.S. 2000. Northwestern Hawaiian Islands derelict fishing net removal project: survey of marine organisms as- 
sociated with net debris. Report submitted to NOAA-NMFS Coral Reef Ecosystem Investigation. 11 pp. 




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Godwin, L.S. L. Harris, A. Charette and R. Moffitt. 2008. The marine invertebrate species associated with the biofouling of 
derelict fishing gear in the Papahanoumokuakea-Marine National Monument. Report submitted to NOAA-NMFS, PIFSC, 
Coral Reef Ecosystem Division. (In Review). 

Godwin, L.S., L.G. Eldredge, and K. Gaut. 2004. The Assessment of Hull Fouling as a Mechanism for the Introduction and 
Dispersal of Marine Alien Species in the Main Hawaiian Island. Final report submitted to the Hawaii Coral Reef Initiative 
Research Program. Bishop Museum Technical Report 28. Contribution 2004-015 to the Hawaii Biological Survey. 

Godwin, S., K.S. Rodgers, and PL. Jokiel. 2006. Reducing potential impact of invasive marine species in the northwest- 
ern Hawaiian islands marine national monument. Report to: Northwest Hawaiian Islands Marine National Monument 
Administration. 

Grigg, R.W. 2003. Invasion of a deep black coral bed by an alien species, Carijoa nisei, off Maui, Hawaii. Coral Reefs 
22:121-122. 

Kahng S.E. 2006. Ecology and ecological impact of an alien octocoral, Carijoa riisei, in Hawaii. PhD thesis, University of 
Hawaii. 

Kilar, J.A. and J. McLachlan. 1986. Ecological Studies of the Alga, Acanthophora spicifera (Vahl) Borg. (Ceramiales: Rho- 
dophyta): Vegetative Fragmentation. J. Exp. Mar. Biol. Ecol. 104: 1-21. 

Littler M.M. and D.S. Littler. 1995. A colonial tunicate smothers corals and coralline algae in the Great Astrolabe Reef, Fiji. 
Coral Reefs 14: 148-149. 

Oda, D. K. and J.D. Parrish. 1982. Ecology of commercial snappers and groupers introduced to Hawaiian reefs. Pp: 
59-67. In Proceedings of the Fourth International Coral Reef Symposium Vol. 1 (E.D. Gomez, C.E. Birkeland, R.W. 
Buddemeier, R.E. Johannes, J.A., Jr. Marsh, and R.T. Tsuda, eds). Quezon City, Philippines: Marine Sciences Center, 
University of the Philippines. 

Preskitt, L. 2002. Acanthophora spicifera (Vahl) Borgesen 1910. Invasive Marine Algae of Hawaii. University of Hawaii 
at Manoa. Fact sheet available from: http://www.hawaii.edu/reefalgae/invasive_algae/rhodo/acanthophora_spicifera.htm 
[Accessed 1 December 2008] 

Pyne, R. 1999. The black striped mussel (Mytilopsis sallei) infestation in Darwin: A clean-up strategy. Ecoports Monogr. 
Ser. No. 19:77-83. 

Randall, J. E. 1987. Introductions of marine fishes to the Hawaiian islands. Bull. Mar. Sci. 41(2): 490-502. 

Randall, J.E., J.L. Earle, T Hayes, C. Pittman, M. Severns, and R.J.F Smith. 1993. Eleven new records and validations 
of shore fishes from the Hawaiian Islands. Pac. Sci. 47(3): 222-239. 

Rodgers, K. and E. Cox. 1999. The rate of spread of the introduced Rhodophytes, Kappaphycus alvarezii (Doty), Kap- 
paphycus striatum Schmitz and Gracilaria salicornia C. ag. and their present distributions in Kane'ohe Bay, Oahu, Hawaii. 
Pacific Science, (53)3: 232-241. 

Russell, D.J. 1987. Introductions and establishment of alien marine algae. Bull Mar Sci, 42: 641-642. 

Russell, D.J. 1992. The ecological invasion of Hawaiian reefs by two marine red algae, Acanthophora spicifera (Vahl) 
Boerg and Hypnea musciformis (Wulfen) J. Ag., and their association with two native species, Laurencia nidifica J. Ag. 
and Hypnea cervicomis J. Ag. ICES mar Sci Symp (Act Symp) 194: 110-125. 

Schumacher, B. D. and J. D. Parrish. 2005. Spatial relationships between an introduced snapper and native goatfishes 
on Hawaiian reefs. Biological Invasions 7: 925-933. 

Shluker, A.D. 2003. State of Hawaii Aquatic Invasive Species Management Plan. The Department of Land and Natural 
Resources, Division of Aquatic Resources. 

Smith, J.E., C.L. Hunter, and CM. Smith. 2002. Distribution and Reproductive Characteristics of Nonindigenous and In- 
vasive Marine Algae in the Hawaiian Islands. Pacific Science 56 (3):299-315. 

Smith J.E., Hunter C.L., Conklin E.J., R. Most, T Sauvage, C. Squair, CM. Smith. 2004. Ecology of the invasive red alga 
Gracilaria salicornia (Rhodophyta) on Oahu, Hawaii. Pac Sci 58: 325-343. 

Southward, A. J., R.S. Burton, S.L. Coles, PR. Dando, R. DeFelice, J. Hoover PE. Parnell, T Yamaguchi, and W.A. New- 
man. 1998. Invasion of Hawaiian shores by an Atlantic barnacle. Mar. Ecol. Prog. Ser. 165: 119-126. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Vargas-Angel, B., L.S. Godwin, J. Asher, and R.E. Brainard. 2008. Invasive didemnid tunicate spreading across coral 
reefs at remote Swains Island, American Samoa. Coral Reefs (In Press). 

Waddell, J.E. and A.M. Clarke (eds.). 2008. The State of Coral Reef Ecosystems of the United States and Pacific Freely 
Associated States: 2008. NOAA Technical Memorandum NOS NCCOS 73. NOAA/NCCOS Center for Coastal Monitoring 
and Assessment's Biogeography Team. Silver Spring, MD. 569 pp. 

Woo, M.M.L. 2000. Ecological impacts interactions of the introduced red alga, Kappaphycus striatum, in Kaneohe Bay, 
Oahu, Masters Thesis, University of Hawaii at Manoa, Honolulu, Hawaii. 

Zabin, C. 2007. A tale of three seas: consistency of natural history traits in a Caribbean-Atlantic barnacle introduced to 
Hawaii. Biological Invasions 9: 523-544 

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waiian Islands. Bishop Museum Occasional Papers 79: 54-58. 



PERSONAL COMMUNICATIONS 

Abbott, I. The University of Hawaii, HI, USA 

Godwin, S. Papahanaumokuakea Marine National Monument Honolulu, HI, USA 
Kahng, S. Hawaii Pacific University, College of Natural Sciences, Waimanalo, HI, USA 
Menza, C. NOAA Biogeography Branch, Silver Spring, MD, USA 



WEBSITES 

University of Hawaii at Manoa • Botany Department and Bishop Museum. Invasive Marine Algae of Hawaii. 2009 
http://hawaii.edu/reefalgae/invasive_algae/index.htm 

German Ministery for Economic Cooperation and Development (BMZ). 2006. http://www.larvalbase.org 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Connectivity and Integrated Ecosystem Studies 

Alan Friedlander 12 , Donald Kobayashi 3 , Brian Bowen 4 , Carl Meyers 4 , Yannis Papastamatiou 4 , Edward DeMartini 3 , 
Frank Parrish 3 , Eric Treml 5 , Carolyn Currin 6 , Anna Hilting 6 , Jonathan Weiss 37 , Chris Kelley 8 , Robert O'Conner 7 , Michael 
Parke 3 , Randy Clark 1 , Rob Toonen 4 and Lisa Wedding 19 

INTRODUCTION 

Population connectivity is the exchange of individuals among geographically separated subpopulations. De- 
fining the scale of connectivity among marine populations and determining the factors driving this exchange 
are critical to our understanding of the population dynamics, genetic structure and biogeography of reef fishes 
(Cowen et al., 2006). Although larvae have the potential for long-distance dispersal, evidence is mounting that 
larval dispersal may be limited and marine subpopulations may be more isolated over smaller spatial scales 
than previously thought (Cowen et al., 2007). The rates, scale and spatial structure of successful exchange, or 
connectivity, among local populations of marine organisms drive population replenishment and, therefore, have 
profound implications for population dynamics and genetics of marine organisms, spatially oriented resource 
management (e.g., marine protected areas) and the spread of invasive species. Despite the importance of this 
issue in understanding population dynamics and effectively managing these species or areas (e.g., Crowder 
et al., 2000; Valles et al., 2001), larval connectivity in the Northwestern Hawaiian Islands (NWHI) is relatively 
unknown. The uniquely endemic fish and other marine faunas of the Hawaiian Archipelago (Hourigan and 
Reese, 1987) and the extreme expression of endemism in the NWHI (DeMartini and Friedlander, 2004) make 
such information critically important for the Hawaiian Archipelago and specifically the Papahanaumokuakea 
Marine National Monument (PMNM). 

LARGE-SCALE POPULATION CONNECTIVITY MODELS FROM OCEAN CURRENTS 

For many marine species, population connectivity is determined largely by ocean currents transporting larvae 
and juveniles between distant patches of suitable habitat. To evaluate the patterns in connectivity throughout 
the Hawaiian Archipelago, a spatially explicit biophysical model was used to simulate coral dispersal between 
reefs spanning the archipelago for three different years (a strong El Nino year- 1997, a strong La Nina year- 
1999, and a neutral year- 2001; Treml et al., 2008). Simulated connectivity was summarized seasonally and 
across years. 

This two-dimensional Eulerian advection-diffusion model of coral dispersal incorporates realistic surface cur- 
rent velocity data and estimates of planktonic larval duration (PLD). In this model, the probability of potential 
dispersal to a reef is the product of: 1) the hydrodynamic arrival probability, 2) larval mortality and 3) the settle- 
ment probability. The spatially explicit hydrodynamic model and resultant arrival probabilities incorporate reef 
topology, ocean current variability and spawning location. 

Summary of Patterns Across Hawaii with Reference to Spatial Data 

Results indicate that the scale of dispersal is on the order of 50-150 km, which is consistent with recent studies 
in the Caribbean (Cowen et al., 2006). On average, the Main Hawaiian Islands (MHI) appear to be consistently 
connected and well mixed at levels above 1/10,000 per season for hypothetical larvae with a PLD of 60 days 
(Figure 9.1). The northwestern most atolls (Kure, Midway, and Pearl and Hermes) are also constantly and 
strongly connected throughout the dispersal scenarios. The entire Hawaiian archipelago appears completely 

1. NOAA/NOS/NCCOS/CCMA Biogeography Branch 

2. The Oceanic Institute 

3. NOAA/NMFS/Pacific Islands Fisheries Science Center, Coral Reef Ecosystem Division 

4. Hawaii Institute of Marine Biology 

5. World Wildlife Fund Fuller Fellow, School of Integrative Biology, University of Queensland, St. Lucia, QLD Australia 

6. NOAA/NOS-Beaufort Lab 

7. Joint Institute of Marine and Atmospheric Research 

8. Hawaii Undersea Research Lab 

9. University of Hawaii at Manoa 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




175'W 



170°W 



1«5*W 



1M"W 



WH 



Figure 9.1. Dispersal pathways in the Hawaiian Archipelago based on Eulerian advection-diffusion models (adapted from 
Treml et a/., 2008). Coral reef habitat is represented by nodes within the graph framework. When larvae from a source 
reef reach a downstream reef site, a dispersal connection is made. This dispersal connection and direction is represented 
by an arrow, or 'edge' within the graph. The thickness of the arrow reflects the strength of connection. Source: Treml, 
unpublished data; map: L. Wedding. 

connected at similar levels for at least one season out of the years modeled, albeit predominately in a north- 
westerly direction. For connectivity via rafting and for those organisms that have a longer PLD or higher sur- 
vival while dispersing, the hydrodynamics around the Hawaiian Islands provide opportunity for dispersal and 
mixing throughout. In addition, long distance larval dispersal from Johnston Atoll to the mid-Hawaii archipelago 
appears to be possible during unique seasons: La Nina, July - September and October - December; neutral 
years, July - September, with the strongest connection in October - December during La Nina years. 



Larval Retention Versus Larval Subsidy 

Metapopulation connectivity in the Hawaiian Archipelago is poorly understood, and this hinders effective man- 
agement and assessment of living marine resources in the region. Pelagic transport was investigated using 
high-resolution ocean current data and computer simulation (Kobayashi, in review). Adjacent strata in the 
archipelago appeared well connected via simulated pelagic larval transport regardless of larval duration, while 
connectivity of more distant strata appear mediated by larval duration (Figures 9.2-9.8). Retention (defined 
as the return of natal propagules) is contrasted with reception or subsidy (the influx of propagules from other 
sources). These two processes appear to be decoupled based on examination of archipelago-wide simula- 
tions. Single-generation and multigeneration effects of connectivity were considered using a simple population 
dynamics model driven by the dispersal kernel probability estimates. The PMNM appears to be largely self- 
sustaining based on these results, with differential input to certain inhabited islands farther southward in the 
archipelago depending on the pelagic larval duration. 

Retention rate (as a fraction of propagules released) ranged from a low of 0.39% at Lanai, to a high of 17.24% 
for the island of Maui (Figure 9.8). When retention and subsidy were pooled to estimate total settlement per 
unit of habitat, settlement ranged from a low of 6,288 settlers per pixel at Kure Atoll to a high of 149,192 set- 
tlers per pixel at Northampton. The high settlement rate at the relatively small Northampton is attributed mostly 
to subsidy. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

The biological significance of the PMNM to the entire Hawaiian Archipelago can be considered from the con- 
nectivity probabilities and the metapopulation analysis. The equilibrium metapopulation composition predicted 
after many generations can be useful in understanding the importance of adjacent or even nonadjacent geo- 
graphic strata. For organisms with short larval duration (15 days), a relatively narrow transitional region in- 
cluding Nihoa, Middle Bank, Niihau and Kauai is composed of settlers from both the PMNM and MHI regions. 
Areas farther north and south have negligible crossover. However, at longer PLDs (90 days), nearly all regions 
throughout the MHI have some component of the settlers derived from the PMNM, whereas most of the PMNM 
is self-seeding until approximately Mokumanamana is reached. 

While the effects of Maro and Gardner can be attributed to their relatively large reproductive output in the 
simulations, other large areas do not contribute similarly to the equilibrium composition, which is a model con- 
sequence of dispersal kernel probabilities operating over many generations. When the effect of habitat size is 
removed by scaling total retention and reception by habitat pixel counts, this yields evidence of a decoupling of 
retention and reception processes. This implies that there is very little, if any, physical (geographic or oceano- 
graphic) relationship between factors which promote effective natal larval retention and factors which promote 
influx of outside larval reception. Settlement and recruitment studies which ignore propagule origins may have 
difficulty in relating observed patterns to oceanographic features for this very reason. Since neither measure 
is a strong proxy for the other, the futility of understanding transport dynamics given the single aggregated 
measure is readily apparent. The need for additional genetics studies and other stock identification markers for 
sourcing of incoming propagules is urgent (e.g., Bernardi et al., 2002; Schultz et al., 2007). 

Clearly since the connectivity measures appear high for adjacent habitats, over evolutionary time the genetic 
connectivity might be more pronounced than inferred here. This could be particularly important at the southern 
boundary of the PMNM, with a protected spawning source able to effectively seed areas to the south over time 
via a "stepping stone" effect, not immediately apparent from examining the pair-wise connectivity values. This 
gradual diffusive process could lead to much more connectivity than that described by a single generation. 



170*E 



173 5 E 



180° 



175'W 



170"W 



165°W 



160 a W 



155°W 



Kure 



'W 



Midway 



Pearl & Hermes 

Maro 



f'*i-'*K;.3."t. ,: *fl. :••_■••-- 

.•JC(V<{* , (. ,fl .f.i. ' 




. N. French Frigate Shoals 
Lisianski f--^_ ." ,yi # \-_^ ±i" *** • • - 

Laysan^V 1 -^' /* — ^Nlhoa J" r%Y ' 

Kauai 



Mokumanamana ' -— S . / Oahu "t ^ 



'*" M.V 
.:4W 



Larval Dispersal (1 yr) 

Percent of total 

Nihoa 

[ J Monument Boundary 



0.0 - 0.0003% 
0.0005% - 0.0008% 
0.0011% - 0,0019% 



0.0022% - 0.0038% 
0.0041% -0.0066% 
0.0068% -0.0101% 



0.0104% -0.0142% 
| 0.0145% ■ 0.0203% 
I 0.0205% - 0.0556% 



Land 



A 



175°E 



180° 



175°W 



170°W 



165°W 



160°W 



155°W 



I 



Figure 9.2. Larval dispersal (45 day PDL) one year after being released from Nihoa Island. Source: Kobayashi, in re- 
view; map: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



170°E 



175°E 



180° 



175°W 



170°W 



165°W 



160°W 



155-W 




Kure 



Midway 



Pearl & Hermes 




Lisianski 



■ L ', 



French Frigate Shoals 



Laysan 

;::"aX;. 

Mokumanamana 



Nillllil 



^v. 



Tu-^rC.= /Maui 



Mo! ok 



■>.:■ 



:;f#tag#^ 



''^a&aV 1 ; 



Larval Dispersal <1 yr) g_ „_„_ 00003% 

Percent of total 

Mokumanamana ■ 0(WO5% " 00008% 

HI Monument Boundary ■ 0.0011% - 0.0016% 



0.0019% -0.003% 
0.0033% - 0.0052% 
0.0055% - 0.0096% 



0.0099%-0.0164% 

0.0167% -0.0249% 

I 0.0252% - 0.2079% 



A 



175°E 



180° 



175°W 



17Q°W 



165°W 



160°W 



155°W 



170°E 



175°E 



175°W 



170°W 



165°W 



160°W 



155*W 



Kure 



Midway 



Pearl & Hermes »JS;«<v 
Maro 






Gardner 




Lisianski 



Laysan 

Mokumanamana 



French Frigate Shoals 
Nihoa 



%, 



-■S^HSL 






. ' '-1- 






uifcC!'..." 
/ Oahu 



,*# 



sir 



Hawaii 



A 



Larval Dispersal (1 yr) 

Percent of total 
French Frigate Shoals 

[ ~^\ Monument Boundary 



| 0.0 - 0.0003% 
0.0005% - 0.0OOB% 
0.0011%-0.0019% 



0.0022% - 0.0036% 
0.0030% ■ 0.0063% 
0.0066% - 0.0099% 



0.0101% -0.0142% 

0.0145% - 0.0214% 

I 0.021 6% -0.0674% 



A 



175°E 



180° 



175°W 



170°W 



165°W 



160°W 



155°W 



Figure 9.3. Larval dispersal (45 day PDL) one year after release from Mokumanamana (top) and French Frigate Shoals 
(bottom). Source: Kobayashi, in review; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



170°E 



173°E 



180° 



175°W 170°W 165°W 160°W 



155°W 




•J&&. 








/Maui 



0.0003% - 0.0005% 



Larval Dispersal (1 yr) 
Percent of total 

Gardner H o.oooe% ■ 0.001 4% 

[ I Monument Boundary I 0.0016% - 0.0025% 



0.0027% - 0.0038% 

o.oo4i%-o.ooe% 

0.0063% - 0.0080% 



0.009% -0.0126% 
| 0.0 129% -0.0 195% 
|0.0197%-0.1364% 



A 



175°E 



180° 



175°W 



170°W 



165°W 



160°W 



155°W 



170'E 



175°E 



180° 



175°W 



170°W 



16S°W 



1S0°W 



155°W 








( I l*v^ Pearl & Hermes ""■J'*^*:. 

Midwa?~\ Vj Maroj^ 
'}".'. -.1 ■ \ . -l^_ Gardner Ej*. ?, 

A 1 \ French Frigate Shoals 

LaysaT — -v " 7^~~ — — ^jNlhoa 

' . j • ^™#£-_ J K ? uai 

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-^vt'-^v'' ' ... •">"■■.": ^,^/Maui 
i&- ,'-\ s "y.rjf £- Molokai-"^ 

v'Sj^T V :*. ' . . ■?/* Hawaii I ,- y 

• ■ :■■■ ■ 






Larval Dispersal 1(1 yr) ^h 0% _ 00u03% 0.0019% - 0.003% o.oos% - o.oi48% I 1 Larid V 

Percent of total I 1 A 

M aro ■ 0.0005% ■ 0.0008% 0.0033% - 0,0052% | | 0.0151% - 0.0247% f\ 

I ] Monument Boundary 0.0011% -0.0016% 0.0056% - 0.0088% | ■ 0.0249% - 0.189% 






— i 1 1 1 1 1 1 





175°E 



180° 



175=W 



170°W 



16S°W 



160°W 



155°W 



Figure 9.4. Larval dispersal (45 day PDL) one year after release from Gardner Pinnacles (top) and Maro Reef (bottom). 
Source: Kobayashi, in review; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



170'E 



175°E 



160° 



175°W 



17CW 



165°W 



1B0*W 



155°W 











Gardnam ■. - 

French Frigate SKoals 



Nihoa 



Mokumanamaria 



auai 



^ 



ahu 



Molokai' 



/Maui 



Hawaii lj--^ 



Larval Dispersal (1 yr) ^g 0% . . 00 03% 

Percent of total 

, o .0005% ■ 0.0008% 
Laysan 

I I Monument Boundary 0.0011% -0.0019% 


0.0022% 

0.0041% 
0.0066% 


0.0038% 0.0099% -0.01 59% I Land 
0.0063% ^B 0.0 162% -0.0263% 
0.0096% ^H 0.0266% - 0.0996% 


N 

A 



175°E 



180° 



175°W 



170°W 



16S°W 



160°W 



155°W 



I 17Q°E 



175°E 



180" 



175°W 



170°W 



165°W 



160'W 



155 a W 






Midway 



Pearl & Hermes 
Maro 



Gardner 




US 



French Frigate Shoals 



Uysan "v 



Nihoa 






>' .' 



Kauai 
, </ ^pahu 

' ; ^j=. /Maui 
Mololiar^JJ 

Hawaii"! .-- > 




Larval Dispersal (1 yr) 

Percent of total 

Lisianski 

I J Monument Boundary 



| 0%- 0.0003% 
| 0.0005% -0.0008% 
0,0011% -0.0016% 



0.0018% -0.003% 
0.0033% -0.0055% 

0.0058% -0,0101% 



0.01 04% -0.017% | ^J Lartd 
| 0.01 73% -0.0285% 
I 0.0293% -0,1614% 



A 



175-E 



180° 



-r 



175°W 



170=W 



165°W 



-r 



160°W 



155°W 



Figure 9.5. Larval dispersal (45 day PDL) one year after release from Laysan Island (top) and Lisianski Island (bottom). 
Source: Kobayashi, in review; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



170*E 



175°E 



180° 



175°W 



17CW 



165°W 



160*W 



155°W 


















I & Hermes •■■!■,'■. 



Midway 







Lisianski 

Laysan 



:. ■■■»:...."■■:. •■■ 



French Frigate Shoals 
Nlhoa 



auai 



; ; -':^' 



Mokumanamana 



J 



ahu 




Molokai 

Hawaii 



Larval Dispersal (1 yr) 

Percent of total 

Pe arl and Hermes 

I ^] Monument Boundary 



| 0%- 0.0003% 
0.0005% - 0.0008% 
0.0011% - 0.0016% 



0.0019% -0.003% 
0.0033% ■ 0.0055% 
0.0058% - 0.0093% 



0.0096% -0.01 59% 

0.0162% - 0.0263% 

I 0.0266% - 0.1014% 



A 



175°E 



180° 



175°W 



170°W 



165°W 



160°W 



155°W 



170*E 



175°E 



180° 



175°W 



170°W 



165°W 



160"W 



155°W 



t*l 



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J* 



ahu 



/Maui 



Molokai 

Hawaii^! ' ' 



Larval Dispersal (1 yr) ■ 
Percent of total 
Midway 

I J Monument Boundary 



| 0%- 0.0003% 
0.0005% ■ 0.0008% 
0.0011% - 0.0018% 



0.0022% - 0.0036% 
0.0038% ■ 0.006% 
0.0063% -0.0093% 



0.0096% -0.0153% [ |_a n( j 

| 0.0156% -0.0271% 
I 0.0274% -0.0701% 



A 



175°E 



180° 



175°W 



170°W 



165°W 



! 

160°W 



155°W 



Figure 9. 6. Larval dispersal (45 day PDL) one year after release from Pearl and Hermes Atoll (top) and Midway Atoll (bot- 
tom). Source: Kobayashi, in review; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



170'E 



175°E 



160° 



175°W 



17CW 



165°W 



150*W 



155°W 






WjWJ 







■-'A i'' 1* .'..■:-■ i 



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Nlhoa 



Mokumanamana 



.!^v^«," 



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, cf Oahu 

Molokai u ;i5 

Hawaii L--' . 



Larval Dispersal (1 yr) 

Percent of total 

Kure 

I J Monument Boundary 



| 0% - 0.0003% 0.001 9% - 0.003% 

0.0005% - 0.0008% 0.0033% ■ 0.0052% 

0.001 1 % - 0.001 6% 0.0055% - 0.009% 



0.0093%-0.0148% 
0.0151% ■ 0.0238% 
0.0241% - 0.0858% 



A 



175°E 



180° 



175°W 



170°W 



16S°W 



160°W 



155°W 



Figure 9. 7. Larval dispersal (45 day PDL) one year after being released from Kure. Source: Kobayashi, in review; map: 
L. Wedding. 



JL 



1 



i 



Hi 



II 



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Figure 9.8: Larval retention for propagules released at each of 10 islands/atolls. The red bars in each graph indicate the 
island or atoll from which the larval propagules were initially released. Source: Kobayashi, unpublished data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




'SSSS'SS/SSSSSS*/ */X/S'/S/*SSsss*S 






W 






<r 



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40 - 
35 - 
30 
25 - 
20 
15 
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Figure 9.8 (continued): Larval retention for propagules released at each of 10 islands/atolls. The red bars in each graph in- 
dicate the island or atoll from which the larval propagules were initially released. Source: Kobayashi, unpublished data. 



Directed Movements of Adult Fishes- Connectivity at the Scale of the Individual 
Acoustic telemetry of giant trevally 
(white ulua, Caranx ignobilis; Figure 
9.9) and jobfish (uku, Aprion virescens; 
Figure 9.9), large-bodied apex preda- 
tors on Hawaiian reefs, revealed each 
to be site attached and home ranging 
(Meyer et al., 2007a,b; Figure 9.10). No 
inter-atoll movements were detected but 
animals were site attached to core activ- 
ity areas where they exhibited diel habi- 
tat shifts and made periodic atoll-wide 

' Figure 9.9. Giant trevally (left) and jobfish (right) are both large, top preda- 

excursions up to IV km. Movements tors in the NWHI coral reef ecosystem. Photos: J. Zamaow and J. Mara- 
to seasonal mating aggregations were gos. 
identified in the summer during specific 
phases of the moon for each species. 





A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Giant Trevally Movement 
A large proportion of giant trevally from 
French Frigate Shoals were caught at 
La Perouse Pinnacle, and these fish 
all showed high detections/day at this 
location, verifying their strong site fidel- 
ity to a core area (Figures 9.11). The 
lower detection/day at East and Tern 
Islands also suggests that these loca- 
tions are on the periphery of the fish's 
home range (none of the fish detected 
at East or Tern were tagged at those lo- 
cations). The large number of fish de- 
tected at Rapture Reef suggests this 
site provides important habitat for giant 
trevally at French Frigate Shoals, as fish 
tagged throughout the atoll made sea- 
sonal excursions to this reef. The arrival 
and departure times of fish were strong- 
ly correlated with each other and in turn 
with the lunar cycle. Coupled with an- 
ecdotal diver observations, the acoustic 
data indicate that Rapture Reef is likely 
a spawning aggregation site for giant 
trevally at French Frigate Shoals. Giant 
trevally tagged at Rapture Reef were 
detected there year round, suggesting 
that their core home range was located 
within the spawning habitat. These fish 
did not make long seasonal movements, 
as their core ranges were within the 
spawning area. The seasonal spawning 
behavior of giant trevally was character- 
ized by daily runs to the spawning loca- 
tions during the lunar spawning cycle. 
They did not shift their core home range 
to the spawning location, as they re- 
turned to their core range (e.g., La Per- 
ouse) after each spawning event. 



VR2 Tracking Data 
Caranx ignobilis 
A Capture locations 
* Rocwwof tocatwfw 

H 

Wa1ar*2Gm ti 
I 1 25 2 5 8 ^ 




-3 




Figure 9.10. Trans-atoll movements of giant trevally at French Frigate 
Shoals (top) and Pearl and Hermes (bottom; Meyer et al., 2007a). Circles 
indicate locations of VR2 receivers, shaded squares indicate giant trevally 
capture sites (numbers within square symbols indicate sites where multiple 
individuals were tagged and released). Lines with arrows indicate most di- 
rect route between giant trevally release and detection locations. 



At Pearl and Hermes Atoll, the great- 
est number of detections of tagged gi- 
ant trevally (for each individual fish) oc- 
curred at the receiver closest to the location where the fish was originally tagged, providing evidence that they 
show strong site fidelity to core areas (Figure 9.12). Giant trevally were detected at receivers at other parts of 
the atoll, suggesting that these areas were on the periphery of the fish's home range. These were most likely 
areas visited during the diel habitat shifts exhibited by almost all individuals. Fish tagged at the Main Chan- 
nel showed greater detections/day at large at the Main Channel receiver, compared to receivers close to fish 
tagged at other parts of the atoll (e.g., northwest corner). The Main Channel is shallow and experiences very 
strong, tidally-driven currents. These strong currents bring animals and materials in and out of the lagoons, 
which appears to make the Main Channel a desirable habitat for apex predators, as suggested by the large 
number of large sharks and teleosts seen at this location. 




so 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Only a limited number of giant trevally were tagged at Midway and Kure Atolls, resulting in few detections and 
no evidence of movement (Figures 9.13 and 9.14). Shallow flats appear to be poor habitat for this species, as 
all receivers located in shallow flats at various atolls recorded very few detections. 









1 

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Figure 9.11. Number of tag detections/days at large for giant trevally tagged at French Frigate Shoals. Source: Fried- 
lander, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 





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Figure 9.11 (continued). Number of tag detections/days at large for giant trevally tagged at French Frigate Shoals. Source: 
Friedlander, unpub. data; maps: L. Wedding. 




W 



J 




Figure 9.12. Number of tag detections/days at large for giant trevally tagged at Pearl and Hermes Atoll. Source: Fried- 
lander, unpub. data; maps: L. Wedding. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




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VR2 Tracking Oata 

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SW 



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Figure 9.12 (continued). Number of tag detections/days at large for giant trevally tagged at Pearl and Hermes. Source: 
Friedlander, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 











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Figure 9.12 (continued). Number of tag detections/days at large for giant trevally tagged at Pearl and Hermes Atoll. 
Source: Friedlander, unpub. data; maps: L. Wedding. 








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Figure 9.13. Number of tag detections/days at large for giant trevally tagged at Midway Atoll. Source: Friedlander, unpub. 
data; maps: L. Wedding. 



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A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 






VR2 Tricking 


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Figure 9.13(continued). Number of tag detections/days at large for giant trevally tagged at Midway Atoll. Friedlander, 
unpub. data; maps: L. Wedding. 









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Figure 9.14. Number of tag detections/days at large for giant trevally tagged at Kure Atoll. Source: Friedlander, unpub. 
data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



VR2 Tracking Dala 
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Figure 9.14 (continued). Number of tag detections/days at large for giant trevally tagged at Kure Atoll. Source: Fried- 
lander, unpub. data; maps: L. Wedding. 

Jobfish Movement 

Movement patterns for jobfish were sim- 
ilar to those observed for giant trevally 
(Figures 9.15). For example, all fish de- 
tected at Rapture Reef were tagged at 
Disappearing Island, located close to 
Rapture Reef (Figure 9.16). However, 
there was no evidence that Rapture 
Reef is a spawning location for jobfish. 
Jobfish tagged along the south coast of 
Pearl and Hermes Atoll were detected 
by receivers on both the southwest and 
southeast tips (Figure 9.17). This sug- 
gests behavior associated with long, 
daily and tidal excursions. 

Overall, jobfish had lower numbers of 
detections/day than giant trevally. This 
may be a function of a key difference in 
their spawning strategy, as well as a ten- 
dency for greater diel movement. Unlike 
giant trevally, jobfish perform complete 
seasonal shifts in their home range, oc- 
cupying separate summer and winter 
core areas. These winter and summer 
locations do not overlap, which is why 
each receiver generally has fewer de- 
tections on an annual basis. However, 
jobfish were occasionally detected in 
their winter or summer location during 
the opposing season, suggesting that 
these seasonal core areas are relatively 
close to each other. 




Figure 9.15. Trans-atoll movements of jobfish at Pearl and Hermes Reef 
with enlarged views of capture areas (insets) showing VR2 receiver loca- 
tions (yellow squares), jobfish capture sites (white circles), jobfish transmit- 
ter codes (white numbers), most direct routes between jobfish release and 
detection locations (dashed red lines). Source: Meyer et ai, 2007b. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Jobfish at Kure atoll also show strong site fidelity to core areas, and fish have been detected at one receiver for 
over three years. Again, however, diel and tidal movements result in jobfish moving over a large area, as ex- 
emplified by the different detection patterns for individual fish seen in Figures 9.18 and 9.19. These fish make 
complete seasonal shifts in habitat as can be seen by the absence of detections during either the summer or 
winter months. The fact that fish were detected, fish that were absent either during the summer or the winter 
months, suggests that the spawning habitats for this species were located at Kure, and that there is more than 
one spawning location at the atoll. 



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Figure 9.16. Number of tag detections/days at large for jobfish tagged at French Frigate Shoals. Source: Friedlander, 
unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



sA 






m 




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Figure 9.17. Number of tag detections/days at large for jobfish tagged at Pearl and Hermes Atoll. Source: Friedlander, 
unpub. data; maps: L. Wedding. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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Figure 9.17 (continued). Number of tag detections/days at large for jobfish tagged at Pearl and Hermes Atoll. Source: 
Friedlander, unpub. data; maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 







•<4 







VR2 Tracking Data 



Qi I 3 




><4 



25.00 

20.00 

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Frigate Point 



602 



Figure 9.18. Number of tag detections/days at large forjobfish tagged at Midway Atoll. Source: Friedlander, unpub. data; 
maps: L. Wedding. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 










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588 604 607 610 



Figure 9.19. Number of tag detections/days at large for jobfish tagged at Kure Atoll. Source: Friedlander, unpub. data; 
maps: L. Wedding. 

Inferring Dispersal and Movement by Tracking Introduced Species 

Eleven species of shallow-water snappers (F. Lutjanidae) and groupers (F. Serranidae) were purposely intro- 
duced to one or more of the main (high) islands of the Hawaiian Archipelago in the late 1950s and early 1960s. 
Of these, three snapper species and one grouper have become established (Randall, 1987). One snapper, 
blueline snapper (Taape or Lutjanus kasmira), and one grouper, Peacock grouper (Roi or Cephalopholis ar- 
gus), are well-established, and have histories of colonization along the island chain that are reasonably well- 
documented. Planktonic stage durations, although unknown for both species, are grossly estimable based on 
congeners elsewhere in the Indo-Pacific. These two species thus represent a unique opportunity to track the 
rate of colonization of introduced species within an oceanic insular ("stepping stone") environment. 

Blueline snapper, if like several other Indo-Pacific congeners, has a planktonic stage duration approximating 
25-47 days and a settlement size greater than 20-30 mm (Leis, 1987), but there is a great deal of geograph- 
ic, seasonal, and other environmental variations in stage duration within and among closely related species 
(Leis, 1993; Victor, 1993). Given these same caveats, Peacock grouper, if a typical member of its genus in 
the subfamily Epinephelinae, settles at a size of about 18 mm (Leis, 1987) and is likely to have a shorter 
pelagic larval stage than blueline snapper. 



A total of about 3,170 blueline snapper were introduced from the Marquesas Islands to Hawaii beginning 
in 1955, including 2,435 released in Kaneohe Bay, Oahu, in 1958 (Oda and Parrish, 1981; Randall, 1987; 
Figure 9.20). The species had colonized the Big Island of Hawaii, 140 nmi downchain of Oahu, by 1960 
(Randall, 1987). Blueline snapper had spread upchain to French Frigate Shoals in the NWHI, 490 nmi from 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Oahu, by sometime between 1977 and 
1982 (Okamoto and Kanenaka, 1984). 
The species was sighted another 330 
nmi farther upchain in the NWHI at Lay- 
san Island (820 nmi from Oahu) in June 
1979 (Parrish et al., 1980; Oda and Par- 
rish, 1981). A few individuals were first 
observed at Midway Atoll, 240 nmi far- 
ther upchain from Laysan island (1,180 
nmi from Oahu), in May-June 1992; 
the species had not been observed on 
similar surveys conducted at Midway in 
1989 and 1991 (Randall et al., 1993). 
These records suggest rates of disper- 
sal of about 18-70 nmi/year for blueline 
snapper subsequent to its introduction 
to Hawaiian waters. This is consistent 
with estimates of realized mean disper- 
sal distance ranging from 33 to 130 km/ 
year from Shanks et al. (2003). 




Figure 9.20. Spread of the introduced blueline snapper (Taape, Lutjanus 
kasmira) throughout the Hawaiian Islands. Source: Sladek Nowlis and 
Friedlander, 2004. 



The dispersal of Peacock grouper fol- 
lowing its introduction to Hawaii is not 
as well documented. However it is clear 
that Peacock grouper has spread less 
extensively than blueline snapper over 
approximately the same time period 
(Figure 9.21). In 1956, a total of 571 
C. argus were introduced from Moorea 
in French Polynesia to Oahu (n=171) 
and to the Kona coast of the Big Island 
(n=400; Randall, 1987). At present, it 
has been documented as far upchain 
as Niihau, 120 nmi from Oahu, where 
it was first observed in November 1978 
(Hobson, 1980). No shallow reef fish 
surveys of the westernmost MHI were 
conducted prior to this time. Peacock 
grouper was absent at French Frigate 
Shoals in 1992 and has been mostly 
absent in annual surveys conducted 
there between 1995 and 2003 (E. De- 
Martini, unpubl. data). Based on this 
meager data, a dispersal rate of >5 
nmi per year is suggested. Although 

pelagic duration estimates are approximate, Peacock grouper- the species with a likely shorter-dura- 
tion pelagic stage- has spread much more slowly through the Hawaiian Archipelago than blueline snap- 
per. Blueline snapper clearly belongs to the long-distance dispersal group (mode greater than 16 km/ 
year); Peacock grouper probably belongs to this group as well, albeit closer to the lower bound. 




Figure 9.21. Spread of the introduced Peacock grouper (Roi, Cephalop- 
holis argus) throughout the Hawaiian Islands. Source: Sladek Nowlis and 
Friedlander, 2004. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Genetic Connectivity Studies 

Ongoing research will determine genetic dispersal among islands and atolls of the NWHI, including both 
invertebrates and reef fishes, using molecular genetic markers to resolve populations and evolutionary parti- 
tions. Preliminary results indicate large differences among taxa in their degree of connectivity throughout the 
archipelago. Some species appear to move around the archipelago with relative ease and show no significant 
population structure in the NWHI and MHI (e.g., reef fish; Schultzetal., 2007; Craig etal., 2007). Other species 
show modest but significant population structure, including the endemic grouper (Rivera et al., 2004), spinner 
dolphins (Andrews et al., 2006) and two damselfishes (Ramon et al., 2008). 




Figure 9.22. A yellowfoot opihi (Cellana sandwicensisj at Kauai. All Hawai- 
ian Cellana spp. are endemic to the archipelago and exhibit a striking popu- 
lation differentiation between the main and northwestern islands. Photo: 
C.E. Bird. 



Opihi, the Hawaiian endemic limpets 
(Cellana exarata; C. sandwicensis, Fig- 
ure 9.22; and C. talcosa), show striking 
population differentiation between the 
MHI and NWHI (Bird et al., 2007). All 
three species of opihi show significant 
differentiation of populations across 
the Hawaiian Archipelago, but the spa- 
tial scales, patterns and magnitudes of 
partitioning differ by almost an order of 
magnitude among species. Preliminary 
data from hermit crabs (Baums et al., in 
prep) indicate variable connectivity in 
this group as well. There is significant 
population differentiation between the 
MHI and NWHI for all three species of 
opihi, and estimates of dispersal (mi- 
grants per generation <3) are so low that 
recruitment from the NWHI would likely 
have negligible impact on depleted MHI 
populations. Even within the MHI, the 
koele (C. talcosa) exhibits such strong 
population differentiation that if the 
Kauai population were depleted, it could 
not recover within our lifetime (Bird et 
al., 2007). 

Kobayashi (2006) recently used a com- 
puter simulation to infer patterns of lar- 
val dispersal between Johnston Atoll 
and the Hawaiian Archipelago. Results 
indicate a "northern corridor" which con- 
nects Johnston Atoll and the central por- 
tion of the NWHI and a "southern corri- 
dor" which connects Johnson Atoll to the 
MHI. Sampling was conducted at John- 
ston Atoll in 2006 to assess connectivity 
between the NWHI and this isolated reef 
habitat. The sea cucumber Holothuria 
atra exhibited low connectivity between 
Oahu and French Frigate Shoals and 
between Oahu and Johnston (Skillings 
et al., in prep; Figure 9.23). In contrast, 

there was no significant difference between samples from French Frigate Shoals and Johnston, supporting 
the northern corridor for dispersal between Johnston and the Hawaiian Archipelago (Figure 9.23). This result 







French Frigate Shoals 








Johnston Atoll 



Figure 9.23. F-statistics demonstrate population genetic separations for the 
sea cucumber Holothuria atra between Oahu (MHI) and French Frigate 
Shoals (NWHI), and between the MHI and Johnston, but high connectivity 
between Johnston and French Frigate Shoals. Source: Skilling et al., in 
prep. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

supports the hypothesis first advanced by Grigg (1981) and Maragos and Jokiel (1986) that Johnston is a po- 
tential gateway that enhances biodiversity in the NWHI. However the alternative hypothesis, that Johnston is 
an outpost of the Hawaiian fauna, remains a viable possibility pending further research. 

Results thus far indicate that population structure across the Hawaiian archipelago does not fit a simple isola- 
tion-by-distance model, and generalizations based on average (geostrophic) oceanographic currents may not 
be warranted (Figures 9.24). Closely-related species with similar ecology and reproductive biology (including 
opihi, hermit crabs, and reef fishes) can have dramatically different patterns of connectivity (Bird et al., 2007; 
Rocha et al., 2007). Together, these results mandate that a suite of invertebrates and fish must be surveyed to 
resolve general trends, and to provide connectivity information pertinent to management of the PMNM. 




Figure 9.24. Apparent shared barrier to dispersal in the Hawaiian Archipelago. Consensus of significant genetic partitions 
among up to 14 marine species across the Hawaiian Archipelago. Locations of apparent restrictions to dispersal are 
marked with yellow bands, and the number of species that share that break out of the total number of species surveyed 
for each location are also given. These results are preliminary but the shared genetic structure among highly divergent 
species thus far is striking. 

Connectivity considerations are particularly important for Hawaiian endemic species. Conservation of Hawai- 
ian endemic species should take into account the consequences of their restricted distribution, including re- 
duced capacity for recovery following depletion. Recently, scientists at the Hawaii Institute of Marine Biology 
have begun to examine population structure in three species of endemic Hawaiian butterflyfishes: the millet- 
seed butterflyfish (Chaetodon miliaris), the bluestripe butterflyfish (C. fremblii) and the pebbled butterflyfish (C. 
multicinctus). Thus far, they have collected and sequenced 170 individuals of bluestripe butterflyfish (Figure 
9.25), 229 milletseed butterflyfish (Figure 9.26) and have made significant progress in collections of pebbled 
butterflyfish (Figure 9.27) throughout the Hawaiian Islands (M.T Craig et al., pers comm.). These species 
perform distinct roles in the coral reef ecosystem and can provide examples of differential connectivity over 
meso-scale distances. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




*st< 0.0001 (P = 0.478, ns) 



Figure 9.25. Haplotype network for the endemic bluestripe butterflyfish. Source: Craig et al., unpub. data. 




Figure 9.26. Haplotype network for of the endemic milletseed butterflyfish. Source: Craig et al., unpub. data. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 




Chaeloc/on muiticinchis 

I Pearl and 1 lermes 
I Laysan 



J Gardner Pinnacles 

| French Frigate Shoals 

| Nihoa 

| Kai 

J Johnsion Atoll 



N = 32 



N = 32 



N = 24 



N = 24 



0>st = 0.00095 (P = 




M=R 



N=g 



N=2 



Figure 9.27. Haplotype network for of the endemic pebbled butterflyfish. Source: Craig et a/., unpub. data. 



ESSENTIAL FISH HABITAT 

Fisheries-habitat links are an important consideration with respect to forms of spatial management such as 
marine protected areas. The composition of suitable habitat within an area can largely dictate fish distribution 
and abundance patterns. The formal concept of essential fish habitat (EFH) was defined with the reauthoriza- 
tion of the U.S. Magnuson-Stevens Fishery Conservation and Management Act in 1996, and refers to habitat 
that is recognized as ecologically important to fisheries resources. Critical fisheries habitats must be identified 
as valued ecosystem components in order to facilitate the formation of ecosystem-based management ac- 
tions. 

Congress defined EFH as "those waters and substrate necessary to fish for spawning, breeding, feeding, or 
growth to maturity" (16 U.S.C. 1802(10)). The EFH guidelines under 50 CFR 600.10 further interpret the EFH 
definition as follows: Waters include aquatic areas and their associated physical, chemical, and biological 
properties that are used by fish and may include aquatic areas historically used by fish where appropriate; 
substrate includes sediment, hard bottom, structures underlying the waters, and associated biological com- 
munities; necessary means the habitat required to support a sustainable fishery and the managed species' 
contribution to a healthy ecosystem; and "spawning, breeding, feeding, or growth to maturity" covers a species' 
full life cycle. 

Analysis was conducted for NWHI bottomfish to determine EFH for these important resource species har- 
vested by Hawaiian-based vessels. The bottomfish fishery has targeted about a half-dozen species of deep- 
slope (generally >75-100 fm) eteline snappers (family Lutjanidae) and one endemic species of epinepheline 
grouper (family Serranidae) out of a total of a dozen common Bottomfish Management Unit Species (WPFMC 
2004; Table 9.1). These species typically inhabitat depth ranges from 100 and 400 m and have been found to 
be associated with certain benthic features, such as high-relief hard-bottom slopes (Kelley, et al., 2006; Kelley 
and Ikehara, 2006; Kelley, 2000). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 9.1. Dominant species in NWHI bottomfish catch and research-based essential fish habitat boundaries (depth in 
meters). Illustrations by Les Hata from Hawaii Divison of Aquatic Resources. 



SCIENTIFIC NAME DEPTH RANGE (m) 




LOCAL NAME 



Ehu 



COMMON NAME 



Red snapper 



Etelis carbunculus 



100-400 




Gindai 



Flower snapper 



Pristipomoides zonatus 



100-350 




Hapuupuu 



Hawaiian grouper 



Epinephelus quernus 



30-300 



Kalekale 



Von Siebold's snapper 



Pristipomoides sieboldii 



50-350 




Lehi 



Onaga 



Reddish snapperfish 



Scarlet snapper 



Apharues rutilans 



Etelis coruscans 



50-250 



100-400 




Opakapaka 



Pink snapper 



Pristipomoides 
filamentosus 



50-300 



EP 



□ 



□3 
-EL □ 



mm^B 



□ o 

I I < 10,000 lbs 
I I 10,000 - 25,000 lbs 
I I 25,000 - 50,000 lbs 
^M > 50,000 lbs 

No Data 
— - Management Boundary 
EEZ 



m 



The NWHI fishery is divided into two 
management zones (Mau, Hoomalu), 
partly in order to distinguish between 
short- and long-duration fishing trips and 
short-duration trips to the closer (to the 
MHI) Mau and more distant (Hoomalu) 
zones, respectively (Figure 9.28). Be- 
tween 1996 and 2004, the Mau zone 
bottomfish catch (Figure 9.29) was 
dominated by shallow-water species 
such as jobfish (39%) and thicklipped 
jack (butaguchi, Pseudocaranx dentex, 
14%), with pink snapper (opakapaka, 
Pristipomoides filamentosus, 13%), Ha- 
waiian grouper (hapuupuu, Epinephelus 
quernus, 13%), and red snapper (ona- 
ga, Etelis coruscans, 8%). In contrast, 
red snapper and pink snapper accounted for 28% and 25% of the Hoomalu catch, respectively, followed by 
Hawaiian grouper (15%). 




\ 



i 



Figure 9.28. Total commercial bottomfish landings from 1996 to 2002. Data 
in several cells can not be shown due to confidentiality concerns. Data: 
DAR; Ehler, 2004. 



The average annual reported landings of bottomfish in the NWHI between 1984 and 2003 were 336,000 lbs 
(SD ± 235,500; NOAA 2006). Of this, the Mau zone averaged 107,130 (SD ± 53,890) or 32% while the aver- 
age catch in the Hoomalu zone averaged 228,730 lbs (SD ± 63,030) or 68% (Figure 9.29). In 2003, the gross 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



reported revenues for the Mau zone 
were $611,000 and $674,000 for the 
Mau and Hoomalu zones, respectively 
(Ehler, 2004). 

In 2003, the number of vessels partici- 
pating in the two zones remained the 
same from the previous year, but there 
were substantial changes in the number 
of fishing trips (NOAA, 2006). In 2003, 
Mau zone trips decreased by 51% re- 
sulting in a 29% drop in landings from 
the previous year. The number of trips 
in the Hoomalu zone increased by 50% 
in 2003, resulting in a 29% increase in 
landings. 

With the initial designation of the NWHI 
Coral Reef Ecosystem Reserve and 
now PMNM, fishing activity in the NWHI 
has been on the decline. Proclamation 
8031 allows commercial fishing by fed- 
erally permitted bottomfish fishery par- 
ticipants that have valid permits until mid-2011 (FR 36443, June 26, 2006). This amounts to a maximum of 
eight permitted bottomfish vessels that fish within the Monument. Significant work was undertaken prior to the 
designation of the Monument in response to previously issued Executive Orders that created the Reserve in 
2000. This fishery operates according to the management regime specified in the Fishery Management Plan 
for Bottomfish and Seamount Groundfish Fisheries in the Western Pacific Region. The management regime 
includes several precautionary measures that minimize potential effects of this fishery. The bottomfishery par- 
ticipants do not operate in the presence of the Hawaiian monk seals and the annual harvest limit for the eight 
vessels is 300,000 lbs. 



j^ Others 

13%_^ 
Onaga S 
8% ^^k 

Hapuupuu 1 y/t 

13% \ yS i 

Opakapaka ■ 
13% 


^k Uku 
\39% 

/ Others 
\ / 9% 

Butaguchi ^k 

14% A ^L 
Onaga fl 

28% m 

Hapuupuu 
15% 


B 

Uku 
12% 

^k Butaguchi 

/ m n% 

1 / Opakapaka 
\^/ 25% 



Figure 9.29. Average species composition (1996-2004) of bottomfish 
catches from the Mau (A) and Hoomalu (B) zones in the NWHI. See text 
for scientific and common names. Source: Kawamoto and Gonzales, 
2005. 



Table 9.2. Optimal bottomfish habitat criteria for NWHI. Source: PIBHMC. 



GIS LAYER SOURCE 



Depth 



Slope 

Backscatter 

Backscatter 



PIBHMC 20 m multibeam data 



Derived from PIBHMC 20 m multibeam data 

R/V AHI 

R/V Kilo Moana 



RANGE 



100-400 m 



The criteria used to delineate poten- 
tial bottomfish habitat in the NWHI was 
based on previous analysis done in the 
MHI (Kelley, 2000; Parke, 2007). Multi- 
beam data sonar provided the GIS layers 
for bottom depth, slope and hardness. 
These factors were used as criteria to 
identify EFH and potential adult habitat for bottomfish (Table 9.2). The depth range found most appropriate for 
this analysis was 100-400 m based on EFH criteria. Areas with slopes greater than 20% were then selected in 
the GIS to further delimit the potential adult habitat areas. Lastly, areas designated as hard bottom based on 
backscatter values were selected for the final potential adult habitat delineation. The range of sonar backscat- 
ter values depended upon the instrument used to collect the data. 



> 20 percent slope 
Hard (>120 m) 
Hard (> 1,000 m) 



EFH and potential adult habitat analy- 
sis was completed for French Frigate 
Shoals, Kure, Maro, and Pearl and 
Hermes because these islands had suf- 
ficient multibeam data. These islands 
currently do not have complete cover- 
age in the depth range designated for 
EFH, so the results of this analysis rep- 
resent bottomfish EFH and potential 



Table 9.3. Area (km 2 ) of EFH based on available multibeam data detailing 
depth (100-400 m) within each island. Source: PIBHMC. 


ISLAND AREA EFH (km 2 ) % OF TOTAL MAPPED AREA 


French Frigate Shoals 


243.93 


23.51 


Kure 


138.79 


31.19 


Maro 


407.97 


30.32 


Pearl and Hermes Atoll 


54.12 


10.98 


Total km 2 


844.81 


24.00 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

adult habitat based on the data available to date. Maps for each island were created for areas that met each of 
the criteria. The areas in these map products that met all three criteria were considered to be "suitable" adult 
bottomfish habitat (Table 9.3; Figure 9.30). 



lee'is'w 166°12"W i66°6 , w 



French 




Frigate 
Shoals J 


pflB^^^ ^^H ->- '• • 






Essential Fish Habitat 

~ ~ J EFH (100-400 m| 

□ ■«« 

Depth (m) 


1 1 Z 4 J\ 

| M ^Ml'l INI 1 1 r\ 


^^^ *^^^^^^flV 





lee'SO'W 166°24'W 



166=18™ lee^iz'w i66°6 , w 



French 
Frigate 
Shoals 




Kure 



Essential Fish Habitat 

| EFH (100-400 in ] 

□ l.,1 
Depth (m) 



A 






178°3tTW 


178°24 , W 


lye'ia'w itb'iz'w 


Kure 




-Mm 




JR^ 










J 




Potential Adult 




Bottomfish Habitat 








H P^nlial Habitat 








Hund 


M ,,; ^P 






Depth (m) 


w~- 4^^ 


^^ ■ 




N 

Bl ~ mw i\ 

1.5 3 6 r\ 

^_ ^^^^Kllnm-^'- 











Maro 



Essential Fish Habitat 

| EFH (100-400m) 



,A 




171°W 


L70"4O'W 170°ZO , W 


Maro 


£/ 


■ # 


Potential Adult 
Bottomfish Habitat 


■■ Potential llriildl 






^La,d 






Depth (m) 






i»o N 
fl i'ii-iirri ■ r\ 









Figure 9.30. Essential fish habitat (yellow) and potential adult bottomfish habitat (red) based on GIS analysis of available 
multibeam data. Maps: L. Wedding. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



176'12'W UB'^W IIWWI 175°54'W 175°48W 175°42'W 



Essential Fish Habitat 

| EFH (100-400 m) 

Depth (m) 




Pearl and 
Hermes 



Potential Adult 
Bottomfish Habitat 



Pearl and 
Hermes 



176°OW l^S^TA/ 175=48™ 175°42 , W 




Figure 9.30 (continued). Essential fish habitat (yellow) and potential adult bottomfish habitat (red) based on GIS analysis 
of available multibeam data. Maps: L. Wedding 



TROPHIC RELATIONSHIPS: STABLE ISOTOPE COMPOSITION OF PRIMARY PRODUCERS 
AND CONSUMER ORGANISMS 



Kure 



Midway 



Pearl & Hermes 




Lisianski 



Laysan 



French Frigate Shoals 
Nihoa 



Mokumanamana 



Kauai 
M Oahu 

Molokai D S5 



Maui 



Locations of Stable 
Isotope Analysis 

1 I Monument Boundary 
{ J Sampling Locations 

P^ Land 



Hawaii 



-^ 



Analysis of the carbon (C) and nitrogen 
(N) stable isotope composition of pri- 
mary producers, benthic invertebrates, 
bony (teleost) fishes and sharks was 
used to assess vertical trophic link- 
ages between primary producers and 
consumer organisms in the NWHI, and 
horizontal trophic linkages between reef 
and pelagic ecosystems. Samples of 
fish, sharks, invertebrates, phytoplank- 
ton and benthic algae were obtained 
during a May 2005 cruise aboard the 
NOAA ship Hiialakai from six locations 
in the NWHI (Figure 9.31). Muscle tis- 
sue was removed from fish (dissection), 
sharks (plugs from tagged animals), and 
invertebrates (dissection). Animal tissue 
was rinsed in distilled water, dried and 
ground prior to stable isotope analy- 
sis. Seawater was prefiltered through a 
200 micron mesh to remove zooplank- 
ton and retain phytoplankton on ashed 
glass fiber filters. Benthic macroalgae were collected by divers, cleaned and rinsed in distilled water, dried and 
ground. Benthic microalgae were collected by divers from surface sediments. Microalgae were separated from 
sediment either by vertical migration through nylon mesh (Currin et al., 2003) or by density centrifugation in 
colloidal silica (Moseman et al., 2004). Algal samples were fumed with concentrated hydrochloric acid to re- 
move carbonates prior to stable isotope analysis. All samples were sent for analysis of 13 C and 15 N composition 
by mass spectrometry at the University of California-Davis. 



A 



Figure 9.31. Locations for stable isotope analysis from cruises in April, May 
and September 2005. Map: L. Wedding. 



The C and N stable isotope composition of algae is a function of ocean chemistry, photosynthesis and growth 
rates, and the specific nitrogen uptake mechanisms of the algae. Typically, phytoplankton have a C isotopic 
signature distinct from benthic algae, and this distinction can be followed through a food web, as animals are 
usually within 0.5%o of the 13 C value of their food. In contrast, algae differ less in their 15 N values, and animals 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

usually have 15 N values that are 2.5 to 4 %o greater than their food, and so N isotopes provide a means to cal- 
culate the number of trophic steps between primary production and a higher level consumer. 

Fish collected from the NWHI were assigned to one of seven trophic groups using diet information in Fried- 
lander and DeMartini (2002) and Parrish and Borland (2004). Invertebrates, which consisted of Hawaiian spiny 
lobster and a single Hawaiian day octopus, were placed in a separate group. All sharks were classified as 
apex predators; see Table 9.4 for list of species collected, trophic group assignments and number of samples 
collected. 



Table 9.4. Trophic group assignments for species collected for stable isotope analysis. Number of samples analyzed (n) 
and common names are also provided. 


TROPHIC GROUP 


GENUS SPECIES 


COMMON NAME 


n 


Herbivores 


Acanthurus olivaceus 


orangeband surgeonfish 


30 


Acanthurus nigrofuscus 


brown surgeonfish 


4 


Acanthurus triostegus 


convict tang 


35 


Acanthurus nigroris 


bluelined surgeonfish 


23 


Stegastes fasciolatus 


Pacific gregory 


5 


Zebrasoma flavescens 


yellow tang 


22 


Corallivore 


Chaetodon lunulatus 


oval butterflyfish 


15 


Zooplanktivores 


Chaetedon milaris 


milletseed butterflyfish 


54 


Dascyllus albisella 


Hawaiian dascyllus 


18 


Myripristis berndti 


big-scale soldierfish 


3 


Myripristis amaena 


brick soldierfish 


3 


Priacanthus meeki 


Hawaiian bigeye 


16 


Invertebrates 


Octopus cyanea 


Hawaiian day octopus 


1 


Panulirus marginatus 


spiny lobster 


42 


Benthic predators 


Lutjanus kasmira 


bluestripe snapper 


11 


Parupeneus porphyreus 


whitesaddled goatfish 


14 


Parupeneus multifasciatus 


manybar goatfish 


37 


Pareupeneus cyclostomus 


yellowsaddle goatfish 


2 


Bodiandus bilunulatus 


Hawaiian hogfish/wrasse 


15 


Chaetodon fremblii 


bluestripe butterflyfish 


30 


Thalassoma ballieui 


blacktail wrasse 


29 


Pelagic predators 


Euthynnus affins 


wavy-backed tuna 


5 


Apex predators 


Caranx melampygus 


blue jack 


19 


Caranx ignobilis 


white jack 


36 


Aprion virescens 


green jobfish 


6 


Carcharhinus amblyrhynchos 


grey reef shark 


6 


Carcharhinus galapagensis 


Galapagos shark 


28 


Galeocerdo cuvier 


tiger shark 


8 


Epinephelus quernus 


Hawaiian grouper 


2 



There was a clear separation in the 13 C signatures of the primary producers in the NWHI system. Phytoplankton 
(Phyto) had an average 13 C value of -23.4%o, consistent with other published values for oceanic phytoplankton. 
Benthic macroalgae (BMA) and microalgae (BMI) were relatively enriched in 13 C, with average values of -18.2 
and -9.5 %o, respectively. There was less separation in the mean 13 N values of benthic algae, which ranged 
from 1.1 to 3.4 %o (Figure 9.32). 

Fish which were a priori placed in the Herbivore category had lower 15 N values than other consumer organisms, 
as expected. However, the offset between the 15 N values of algae and several members of the Herbivore group 
was higher than the expected 2 to 4/mil, suggesting that either some of the fish designated as herbivores are in 
fact omnivores, or that the algal N values obtained during the May 2005 cruise were more depleted than algal 
values earlier in the season. This latter point can reasonably explain the observed data, as the isotope com- 
position of fish tissue turns over much more slowly than the isotopic composition of the faster growing algae. 
There was also a significant range in the C values within the Herbivore group, with yellow tang in particular 





A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



closest to the phytoplankton endmem- 
ber, and the brown surgeonfish closest 
to the benthic microalgal endmember. 
Both yellow tang and brown surgeonfish 
are browsers of macroalgae but isotopic 
differences suggest there may be finer 
scale variations in diet (Jones, 1968). 



Fish designated as Zooplanktivores and 
Corallivores, as well as lobster and oc- 
topus, had 15 N values of approximately 
8.25%o, consistent with feeding at two 
trophic levels above primary producers 
with an average 15 N trophic fractionation 
value of 2%o. The Zooplanktivores had 
the lowest C isotope signature, sug- 
gesting a greater contribution of phy- 
toplankton to their food web, although 
a significant portion of benthic produc- 
tion was also utilized by this group. In 
contrast, the Corallivore had a more en- 
riched C isotope signature, consistent 
with a greater contribution of coral and 
benthic algae to its food web. 

Fish designated as Benthic Predators 
exhibited approximately a 3%o range 
in both C and N isotope signatures. 
Benthic Predators with 15 N values >10 
(blacktail wrasse) may be feeding 2.5 
to 3.0 trophic levels above the primary 
producers. The Benthic Predators with 
the most enriched 13 C values were the 
whitesaddle goatfish and the Hawaiian 
hogfish, 



14 



12 



10 



Z 



^ Tiger shark 



Hawaiian Grouper 



* 






whitesaddled 
goatfish 



brick ___ 
soldierfish 

yellow 
tang 



V* 



V 



-H3p 



^y^ 

O o r 



brown 
surgeonfish 







! 



o 


Herbivores 


■ 


Zooplanktivores 


▼ 


Corallivores 


▲ 


Benthic predators 


• 


Apex predators 





Pelagic predator 


■ 


BMA 


■ 


BMI 


■ 


Phyto 



+ 



-25 



-20 



-15 



-10 



§13c (%„) 



Figure 9.32. Dual isotope plot of consumer and producer groups from 
NWHI. Each symbol represents the mean ± one standard error of the 15 N or 
13 C value for a species offish, shark or invertebrate. Species list, number of 
samples and trophic group designations are as in Table 9.4; arrows point to 
species referred to in the text. 



The 13 C values of fish designated as Pelagic Predators were relatively depleted in 13 C, suggesting that phyto- 
plankton did contribute substantially to the food webs supporting these fish. The 15 N values of Pelagic Preda- 
tors averaged 9.3%o, which is very similar to the average trophic level of the Benthic Predator group, and 
consistent with an organism feeding two to three levels above the primary producers. 

The 15 N values of fish and sharks designated as Apex Predators overlapped with the 15 N values of the Pelagic 
Predator (wavy-backed tuna) or Benthic Predator (blacktail wrasse). The exception is the enriched 15 N value of 
12.1 for the tiger shark, which puts it nearly a full trophic level above other predators in the NWHI ecosystem. 
This is consistent with marine mammals, sharks, birds and other upper trophic level prey comprising a larger 
portion of the tiger shark diet than that of Galapagos and grey reef sharks (Papastamatiou etal., 2006). The 13 C 
values of several of the Apex Predator group were enriched in 13 C compared to other predators in the system. 
In particular, it appears that grey reef sharks, Galapagos sharks, giant trevally, and tiger sharks are obtaining 
the bulk of their C from a benthic-based food web. 



The relative contribution of benthic primary production to the food webs supporting bony fish, shark and in- 
vertebrate production can be estimated by comparing the stable isotopic composition of these groups with 
values that would be expected from a prescribed food web. In Figure 9.33, the mean isotope values of each 
of the trophic groups described in Table 9.4 are displayed. The black dotted lines in the figure represent the 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



15 
14 
13 
12 
11 
10 
9 
8 



4TL 



3TL 



£ 



If) 

to 



2TL 



iO 



o 


Herbivore 


▼ 


Corallivore 


■ 


Zooplanktivore 


o 


Invertebrate 


▲ 


Benthic Predator 





Pelagic Predator 


• 


Apex Predator 




Trophic Levels (1 - 4) 




assuming food base 




of 33% Phyto, 33% BMA 




and 33% BM1 



o 



1TL 



* 



* 



BMI 



expected graphical position of a con- 
sumer group feeding at the first Trophic 
Level (1 TL) through the fourth Trophic 
Level (4 TL). The position of these black 
lines is based on the assumption that 
there is a 2 - 4%o increase in 15 N values 
per trophic step, and a 0.5%o increase 
in 13 C values per trophic step, and that 
the food web is based on equal parts 
phytoplankton, benthic microalgae, and 
benthic macroalgae. In terms of trophic 
level of the various groups, the figure 
clearly illustrates the discrepancy be- 
tween the 15N values of the primary 
producers as measured in May 2005 
and the herbivores that are presumably 
grazing on them. As noted previously, 
this could be due to a short-term de- 
crease in the 15 N values of the primary 
producers, and the longer-term average 
value of the primary producers may be 
closer to 3.5 - 4.5%o, which would re- 
sult in the observed herbivore 15 N val- 
ues. Alternatively, it may be that some 
members of the group designated as 
'Herbivores' are in fact. Figure 9.33 also 
clearly illustrates that the groups desig- 
nated as Zooplanktivores, Corallivores 
and Invertebrates (lobster) are feeding 
a full trophic level above the Herbivores, 
and that the Benthic and Pelagic Preda- 
tor groups are feeding about one-half 
trophic level above that position. Apex 
Predators (sharks and jacks) are feed- 
ing nearly a full trophic level above the 
Invertebrate/Corallivore level, and about 
one-half trophic level above the Benthic 
and Pelagic Predators. As noted previ- 
ously, and illustrated in Figure 9.32, tiger sharks are an exception and are feeding a full trophic level above the 
Benthic Predator group. The figure also reveals that all groups other than Pelagic Predators, Zooplanktivores 
and Corallivores fall roughly where they would be expected to fall if phytoplankton represented approximately 
33% of the base of their food web, with the remaining portion deriving from equal parts benthic microalgae and 
benthic macroalgae. The position of the exceptions indicate that phytoplankton represent a greater proportion 
of the food web support for Pelagic Predators and Zooplanktivores, and that phytoplankton represent less 
than a third of the food web support for Corallivores. Taken together, these results from analysis of the stable 
isotope composition of primary producers and consumers from the NWHI are remarkably consistent with the 
Ecopath model estimates of the food web supporting fishery production in the NWHI (see next section). Both 
approaches indicate that benthic algae provide the majority of trophic support for apex predators, and that the 
entire system consists of a relatively short (three to four trophic levels above primary production) food chain. 



Phyto 



BMA 

i 



-24 -22 -20 -18 -16 -14 

5 13 C (%o) 



-12 -10 -8 



Figure 9.33. Dual isotope plot of mean isotope values of primary producers 
and each of the trophic groups described in Table 9.4. The black dotted 
lines in the figure represent the expected graphical positions of a consumer 
group feeding at the first Trophic Level (1 TL) through the fourth Trophic 
Level (4 TL), assuming a food web based on equal parts of each of the 
three primary producers (phytoplankton, benthic macroalgae, benthic mi- 
croalgae). Further details on assumptions are in the text. 





Figure 9.34. The coral reefs of the NWHI are a very diverse and unique 
ecosystem, providing habitat for a wide range of marine life. Photo: J. Mara- 
gos. 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

FOOD WEB MODELS 

The coral reefs of the NWHI represent a 
diverse marine ecosystem that provides 
habitat to a number of organisms (Fig- 
ure 9.34). In the mid to late 1970s, doz- 
ens of scientists participated in a large, 
multi-year field study program at French 
Frigate Shoals to describe and better 
understand this ecosystem (Grigg et al., 
2008). These efforts yielded Ecopath, a 
simulation program designed to model 
the flow of energy throughout the sys- 
tem. Ecopath works by creating a snap- 
shot of the ecosystem and the feeding 
relationships between species within 
that ecosystem. The trophically linked 
components consist of a single spe- 
cies, or a group of species represent- 
ing ecological levels. For each species 
group, biomass, production/biomass ra- 
tio (or total mortality), consumption/bio- 
mass ratio and ecotrophic efficiency are 
measured (Polovina, 1984). Ecosim, a 
new dynamic modeling program based 
on the original Ecopath model, is now 
available at (http://www.ecopath.org). 

Ecopath was first applied to data col- 
lected at French Frigate Shoals during 
the late 1970s (Figure 9.35). The eco- 
system was divided into 12 species 
groups with sharks, jacks, monk seals, 
sea birds and tuna at the top trophic lev- 
el, reef fishes at the center, and benthic 
algae, responsible for 90% of the pro- 
ductivity, at the bottom (Polovina, 1984). 
The large reef fishes group was further 
divided into four feeding guilds, result- 
ing in an ecosystem spanning almost 
five trophic levels with sharks, jacks and 
piscivorous reef fish representing the 
top predators (Polovina, 1984). With the 
exception of limited handline fishing for 
snappers, the NWHI are not fished and 

experience relatively few, severe local anthropogenic threats (although sea level rise, acidification, and the 
warming/bleaching and loss of coral habitat will likely become a major human agent of change at basin and 
global scales later in this century). Because the NWHI presently has few severe local threats, the Ecopath 
model provides a picture of an increasingly rare coral reef ecosystem dominated by an abundance of apex 
predators. 

The Ecosim was used to simulate changes in ecosystem dynamics over time in response to top-down or 
bottom-up forcing (Christensen and Walters, 2004) which was modelled by assuming 30 years of high benthic 
primary productivity, followed by 30 years of low benthic primary productivity (Grigg et al., 2008). Significant 




ALGAE 
2.0x10 s 
'= 2.5x10 s 



Figure 9.35. Illustration of the Ecopath Model for the food web at French 
Frigate Shoals. The trophic pathway annual production (P), and mean an- 
nual biomass (B; kg/km 2 ) is given for 12 species groups based on an area 
of 1,200 km 2 . Source: Polovina, 1984. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

temporal lags, varying by as much as a decade, were observed in the responses of the various trophic guilds 
both under an increase and a decrease in benthic productivity (Grigg et al., 2008). Planktivorous reef fish 
trended downward when benthic productivity was high due to the increase in predatory species (e.g., jacks). 
This was the case even when prey plankton was unchanged. When benthic productivity was changed from 
high to low there was an immediate sharp increase followed by a decline in benthic carnivorous reef fishes. 
The reef fishes quickly increased in abundance in response to higher prey availability, but five years later as 
their predators increased, their abundance declined (Grigg et al., 2008). Even with the more complex Ecosim 
model, it is important to note that ecosystem dynamics are more complicated than the model provides and are 
not always consistent with model forcing. 

In the last 10 years the Ecosim model was revised using updated parameters and a reference biomass based 
on surveys of benthic/demersal fish taxa that exhibited habitat fidelity (Parrish, unpub data). Field surveys were 
spatially stratified by the region's primary habitat types in order to make the model more accurate (Grigg et al., 
2008). The surveyed fish communities occupied the central portion of the ecosystem food web and were used 
to project a minimum biomass for the lower guilds, as well as a theoretical maximum value for the top level 
transient predators that preyed on the fish (Grigg et al., 2008). Work is now underway to validate the model 
with the best field estimates of population size, body size distributions, and size-specific food and feeding hab- 
its for the endangered Hawaiian monk seal, a top level predator in the NWHI. Numbers and body condition of 
the seals have been closely monitored for the last two decades and foraging studies indicate a diet of primarily 
benthic/demersal fish (Goodman-Lowe, 1998; Parrish et al., 2000, 2002 and 2005). Successful validation of 
the model using monk seals will depend on knowing the boundaries of the seal foraging activity and the relative 
composition of the diet (Grigg et al., 2008). Once initial validation efforts are complete, the dynamic simulation 
phase using Ecosim (Figure 9.36) will begin with the goal of forecasting and hindcasting situations to illustrate 
how the system might react to both natural and anthropogenic stressors. 



^Biomass/Original Biomass 




Turtles 

BankPisc 

RFPisc 

Heterotrophic Benthos 

Reef Sharks 

Cephalopods 

Bank Jacks 

RFBC 

Bank Sharks 

Reef Jacks 

RF Herbiv 

Bank Herb 

Macro Heterotrophs 

Tiger Sharks 

Detritus 

BankBC 

Benthic Algae 



Figure 9.36. Capture of the Ecosim software output for the Ecopath model at French Frigate Shoals. Display shows the 
response of ecosystem component to a 50% reduction in benthic algae. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

EXISTING DATA GAPS 

To understand passive transport, there is a need for basic information on spatial and temporal patterns of water 
movement, quality and characteristics within the NWHI at a range of scales to determine the general patterns 
of passive transport for nutrients and living resources. Building on an understanding of oceanographic pro- 
cesses, specific research needs and opportunities include efforts to: 

Determine the transport pathways and patterns for the larvae of key organisms; 

Identify the sources and sinks of larval dispersal for key organisms; 

Define the sources and patterns of primary productivity resulting from upwelling sites and occurrences 
and nutrient input to the NWHI; and 

Undertake applied research into the design of protected areas in support of ecosystem resilience 
based on passive transports processes, patterns and pathways. 

Overall there is a need for systematic information on the active transport and movement of biota into, out of 
and within the NWHI. This work can be extended to important applications such as stock identification, popula- 
tion dynamics and species interactions. All of these efforts should be undertaken in a way that contributes to 
the development of models that can predict movement patterns at multiple spatial scales to address questions 
of connectivity, including the linkages between the NWHI and the MHI. 

Specific opportunities include research to improve the understanding of: 

What are the important species that have regular or episodic, active movements or migrations into and 
out of the NWHI and MHI? 

What life stages of these species are involved in the active movements? 

What are the important habitats for different life stages of these species that move among the reefs 
within the NWHI and between the NWHI and MHI? 

What are the effects of extreme events and anthropogenic stressors on movements and migrations? 

Which habitats are at risk from climate change and other forces (e.g., sea turtles and their nesting 
beach habitat)? 

As the understanding of most of the species and populations in the NWHI is at the most basic level (e.g., 
identification of species and groups), genetic studies have the capability to enhance the understanding of the 
ecosystem, including distribution, dispersion rates, and connectivity or isolation among plant and animal popu- 
lations in the NWHI. Specific research opportunities include: 

Characterizing the genetic structure of key species and populations; 

Determining genetically distinct subpopulations of flora or fauna between the MHI and the NWHI; 

Determining the value of selected species in the NWHI for repopulating MHI populations that are over 
exploited or subject to major impacts; 

Applying genetic techniques to key populations across the stress gradient of the archipelago to detect 
pools of individuals with a genetic makeup that keeps them from being filtered out by the environmental 
stressors; 

Studying individual species' response to natural and anthropogenic stress (determining the coral spe- 
cies that are more heat tolerant and can withstand coral bleaching); 

Identifying key species that may be at risk from the genetic influence of invasive species. Identifying 
pilot taxa to serve as proxies for ecosystem genetic connectivity; and 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Because management decision making will be improved by knowledge of many specific factors influ- 
encing ecosystem resilience, information is needed on resilience pathways, such as: acclimation to 
stress, adaptation to stress, the role of the environment and the role of the community. Specific ex- 
amples of research opportunities include activities to determine: 

The key aspects that affect ecosystem stability and resilience (e.g., rates of energy flow, ocean- 

ographic conditions, nutrient levels and recruitment); 

The degree to which natural variability in an ecosystem may determine its capacity for resil- 
ience; 

How ecosystem acclimation to change varies among taxa and in relation to survival and the 

ability to effectively reproduce; 

How genetic makeup enhances the ability of taxa to recover from some kinds of stress; 

Which environmental conditions, e.g., temperature, flow, geomorphology, have a mitigating in- 
fluence on survival in a changed environment; 

The extent to which the reduction or expansion of one or more species or functional groups 

results in top down predation or an increase in bottom up production; 

How the rebound of an ecosystem depends on maintaining established pathways of energy flow 

which provide the system a stable means of recovery rather than risk a transition to a different 

state of equilibrium; 

The extent to which reducing fish populations of the ecosystem undermine or realign energy 

flow and trophic stability; and 

Whether self seeding systems are resilient. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

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ies of the Western Pacific region: 2002 Annual Report. Western Pacific Regional Fisheries Management Council, Hono- 
lulu, Hawaii. 



PERSONAL COMMUNICATIONS 

Craig, M.T. Hawaii Institute of Marine Biology, Kaneohe, HI, USA. 

WEBSITES 

University of British Columbia (UBC), Fisheries Centre, http://www.ecopath.org 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Management Concerns and Responsibilities 

Kaylene E. Keller 1 , Angela D. Anders 2 , Ann Mooney 1 , Randall Kosaki 1 , Malia Chow 1 and Mark Monaco 3 



INTRODUCTION 

Increasing pressures on the world's ocean resources in the recent decades has heightened the need for pro- 
tecting marine resources. Marine Protected Areas (MPA) are an essential tool for achieving marine ecosys- 
tem-based management. MPAs in the Pacific, such as Papahanaumokuakea Marine National Monument, the 
Australia's Great Barrier Reef Marine Park and the Phoenix Islands Protected Area in Kiribati lead the world in 
protecting large-scale marine ecosystems. Each of these MPAs conserves vast contiguous areas of ecosys- 
tems ranging from shallow water coral reefs to deep water communities. 

The Papahanaumokuakea Marine National Monument (Monument) is one of the largest and most unique 
MPAs in the world. The Monument contains relatively pristine ecosystems and cultural resources minimally af- 
fected by human activities. In seeking to preserve and protect these attributes, Monument managers identified 
the following mission: "Carry out seamless integrated management to ensure ecological integrity and achieve 
strong, long-term protection and perpetuation of the NWHI ecosystems, Native Hawaiian culture, and heritage 
resources for the current and future generations" (Papahanaumokuakea Marine National Monument, 2008). 

Striving to achieve this far-reaching mission creates unique opportunities and challenges for Monument man- 
agers. These opportunities include the potential to manage complete ecosystems with few anthropogenic 
inputs and working toward restoring components of the ecosystems that have been modified. Challenges re- 
volve around the remote and vast nature of the Monument, and include threats local to global and internal and 
external to the Monument. This chapter focuses on management of the Monument. This includes: 

• Management structure; 

• Management of protected marine species within the Monument; 

• Management of greatest potential threats to the marine resources across the region; and 

• Management of human activities. 



BACKGROUND 

Management Structure 

On June 15, 2006, President George W. Bush issued Presidential Proclamation 8031 (Proclamation) estab- 
lishing the NWHI Marine National Monument under the authority of the Antiquities Act of 1906 (16 U.S.C. 431). 
It was subsequently renamed the Papahanaumokuakea Marine National Monument. The Monument includes 
a number of preexisting federal conservation areas: the NWHI Coral Reef Ecosystem Reserve, managed by 
the Department of Commerce through the National Oceanographic and Atmospheric Administration (NOAA) 
Office of National Marine Sanctuaries; Midway Atoll National Wildlife Refuge, Hawaiian Islands National Wild- 
life Refuge, and Battle of Midway National Memorial, managed by the Department of the Interior through the 
United States Fish and Wildlife Service (USFWS). These areas remain in place within the Monument, subject 
to their applicable laws and regulations in addition to the provisions of the Proclamation. 

The NWHI also include state of Hawaii lands and waters, managed by the Hawaii Department of Land and 
Natural Resources (DLNR) as the NWHI Marine Refuge and the State Seabird Sanctuary at Kure Atoll. These 
areas also remain in place and are subject to their applicable laws and regulations. 



1. NOAA/NOS/ONMS/Papahanaumokuakea Marine National Monument 

2. Clancy Environmental Consultants, Inc. 

3. NOAA/NOS/NCCOS/CCMA Biogeography Branch 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

The organizational structure for the Monument consists of: 

• Three Co-Trustees-- Department of Commerce, Department of the Interior and the State of Hawaii- re- 
sponsible for the management of the Monument. The Co-Trustee agencies have developed a joint man- 
agement plan that will guide management of the Monument for the next 15 years; 

• A Senior Executive Board composed of a designated senior policy official from each Co-Trustee agency 
that is directly responsible for providing oversight and guidance for management of the Monument; 

• A Monument Management Board composed of representatives from the federal and state agency offices 
that carry out the day-to-day management and coordination of Monument activities; and 

• An Interagency Coordinating Committee representing other state and federal agencies as appropriate to 
assist in the implementation of Monument management activities. 

Management Zones 

Monument regulations define three types of marine zones within the Monument (Figure 10.1): 

1. Special Preservation Areas: These are discrete, biologically important areas of the Monument where re- 
source harvest and almost all forms of discharge are prohibited; 

2. Ecological Reserves: These areas consist of contiguous, diverse habitats that provide natural spawn- 
ing, nursery, and permanent residence areas. Resource extraction is highly restricted within Ecological 
Reserves; and 

3. Midway Atoll Special Management Area (SMA): Recreational activities in the Monument are restricted to 
the Midway Atoll SMA. 

Zoning not only provides protection to highly sensitive habitats, it also protects the ecological linkages between 
these habitats. Each zone addresses a number of factors including the protection of habitat and foraging areas 
of threatened and endangered species; the inclusion of a representative range of the diverse array of marine 



Papahanaumokuakea 
Marine National Monument 



Legend 



\ Marine National Monument Boundary 
_] 100 Fathom Contours 
Special Preservation Area 

IGOfm: KureAloll. Pearl and Hermes Atoll 

50fm: Layssn Island 

25lrn Ltennskl Island. Mara Reel. Gardner pinnacles. Necker Island 

3nm: Ninoa 

Ranch frigate Shoals Boundary Ceorainales: 

(166* 45' W. 24* 10" Nj {IBS' 35' W. 24* Hh 

(166* 45' W. 24' 2' NJ (166* Hf W. 23' 41' U) 

(166* 5S 1 W. 24' 2 N) (165* 35~ W, 23' 30" Ni 

' Ecological Reserves 
^J Existing Commercial Fishing Area Phased Out by June 2011 

Managed as Ecological Reserve fbffctw'ig phase-out 

i Special Management Area 

Boundary extends 1?nm tnjfn land 

1"^ j Emergent Land Features 




Source Data pfcwided by HOAA. gale c* Hawaii and SSRa 



— I 

:c: i. 



Figure 10.1. Map of the Papahanaumokuakea Marine National Monument and zones. Map: PMNM. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

habitats, including shallow coral reef environments, as well as deepwater slopes, banks and seamounts; and 
finally, the minimization of risks associated with specific activities such as fishing and recreational activities. As 
of June 2011 all commercial fishing will be prohibited within the Monument. 



In addition to the designation of the Mon- 
ument management zones, in 2007 the 
Monument was designated "in principle" 
as a Particularly Sensitive Sea Area 
(PSSA) by the International Maritime Or- 
ganization (IMO), a Specialized Agency 
of the United Nations (Figure 10.2). The 
designation puts into effect internation- 
ally recognized measures designed to 
protect marine resources of ecological 
or cultural significance from damage by 
ships while helping keep mariners safe. 
PSSA designation augments domestic 
protective measures by alerting inter- 
national mariners to exercise extreme 
caution when navigating through the 
area. A U.S. proposal for PSSA desig- 
nation was submitted in April 2007 for 
consideration at the IMO's Marine Envi- 




I Papahanaumokuakea Marine National Monument 
and Particularly Sensitive Sea Area 



| Ship Reporting Area 

Aieas to be Avoided 

Data SOuit&: NOAA. NGDC Aritf ESRI<& 



400 Kilometers 



Not ttf tie used fm ftSavrtjitt/Qti 



— I — 

200 



A 



Figure 10.2. PSSA, Areas to be Avoided (ATBA) and Ship Reporting Bound- 
aries around the Papahanaumokuakea Marine National Monument. 



ronment Protection Committee meeting with the final designation made in April 2008. PSSA designation has 
been granted to only 10 marine areas globally, including the marine areas around the Florida Keys, the Great 
Barrier Reef and the Galapagos Islands. The PSSA area is coterminous with the Monument boundary. 

In addition to alerting international mariners to exercise extreme caution when in the area, as part of the PSSA 
designation process, the IMO's Maritime Safety Committee adopted the U.S proposals for the associated pro- 
tective measures (APMs) of: (1) the expansion and amendment of the six existing recommendatory Areas to 
be Avoided (ATBAs) in the area, which would enlarge the class of vessels to which they apply and augment the 
geographic scope of these areas, as well as add new ATBAs around Kure and Midway atolls; and (2) the es- 
tablishment of a ship reporting system for vessels transiting the Monument, which is mandatory for ships 300 
gross tons or greater entering or departing a U.S. port or place and recommendatory for other ships. These 
APMs were implemented in May 2008. 



WORLD HERITAGE NOMINATION 

The unique habitats and ecosystems within the Monument are of great importance to local, regional and global 
marine biodiversity (Figure 10.3). The Monument contains some of the world's most significant marine and 
terrestrial ecosystems and areas of cultural significance, and is one of the world's largest protected marine 
areas. It also serves as an example of ongoing geological processes and biological evolution. The volcanic 
rocks, large atolls of sand and coral, and islets surrounded by reefs provide unique habitats for endemic and 
rare species of animals and plants. These features are of universal value from scientific, conservation, cultural 
and aesthetic perspectives. This relatively pristine region contrasts sharply with most insular and marine eco- 
systems, which are more severely affected by human activities and populations around the world. 



In January 2008, Papahanaumokuakea was selected by the Secretary of the Interior to be included as a can- 
didate for the U.S. Tentative List for nomination as a World Heritage mixed site due to its exceptional natural 
and cultural importance. World Heritage is the designation for places on earth that are of outstanding univer- 
sal value to humanity and as such, have been inscribed on the World Heritage List to be protected for future 
generations to appreciate and enjoy. In early 2009, the U.S. put forth a full nomination package to the World 
Heritage Centre to have the Monument added to the World Heritage List. The Monument was recommended 





A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

for consideration as a World Heritage 
Mixed Site. The reasons for listing for 
the Monument for natural values in- 
clude: 

• The string of islands comprises 
a classic, important and unparal- 
leled example of later stages of 
island and atoll evolution. The ar- 
chipelago has provided some of 
the most compelling confirmation 
of current theories of global plate 
tectonic movements; 

• Papahanaumokuakea is a spec- 
tacular example of evolution in iso- 
lation, which results in enhanced 
speciation and a phenomenally 
high degree of endemism in both 
marine and terrestrial flora and 
fauna. The coral reef ecosystems 
of Papahanaumokuakea also represent on of the worlds's last apex predator dominated ecosystems, a 
community structure characteristic of coral reefs prior to significant human exploitation; and 

• The region is home to, and a crucial refuge for, many endangered, threatened, and endemic species, in- 
cluding critically endangered marine mammal, bird, and plant species for whom it is the last or only refuge 
anywhere on earth. Papahanaumokuakea is also the largest tropical seabird rookery in the world. 

Remote, uninhabited and relatively pristine in comparison to other marine ecosystems in the world, the Monu- 
ment has the potential to serve as one of the few reference sights for monitoring and deciphering short-term 
and long-term responses to local, regional, and global environmental and anthropogenic stressors. The Monu- 
ment is one of the few regions on Earth where monitoring and research activities can be conducted in the 
virtual absence of local human habitation. In comparison, most reef systems in the coastal regions of the 
world are adjacent to human population centers, where vessel traffic, overharvesting, sedimentation, habitat 
destruction, and other human actions have altered the terrestrial and adjacent marine environments. Ongoing 
research, monitoring, habitat restoration and conservation management of the insular and marine ecosystems 
in the NWHI will continue to provide significant insights that will benefit management interventions not only for 
the NWHI, but for insular and marine ecosystems around the world. 



Figure 10.3. The variety of ecosystems within the Monument have been 
recognized for their uniqueness and importance to global marine biodiver- 
sity. Photos: J. Watt. 



MANAGEMENT OF PROTECTED SPECIES 

The NWHI provides habitat for a wide variety of species including species specifically protected by federal acts 
and state statutes. Three federal acts, as well as multiple state statutes, provide protections for specific spe- 
cies in the NWHI. The federal acts are the Endangered Species Act (ESA), the Marine Mammal Protection Act 
(MMPA) and the Migratory Bird Treaty Act (MBTA). The ESA of 1973 provides for the conservation of species 
at risk of extinction throughout all or a significant portion of their range, and the conservation of the ecosystems 
on which they depend. The state of Hawaii has adopted specific criteria for indigenous species to be listed as 
threatened or endangered, as codified in chapter 195D-4, Hawaii Revised Statutes (HRS), as well as chapter 
183D, HRS Wildlife, and chapter 125, Wildlife Sanctuaries, Hawaii Administrative Rules Title 13. The MMPA 
provides protection and conservation of all marine mammals whether or not listed under the ESA. The MBTA is 
a domestic law that implements the United States' commitment to four international conventions (with Canada, 
Japan, Mexico and Russia) for the protection of shared migratory bird resources. All migratory birds and their 
parts (including eggs, nests and feathers) are fully protected. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

The Monument provides habitat for many protected marine species including the Hawaiian monk seal, five 
cetacean species, five marine turtles and five bird species (Figure 10.4). 




Figure 10.4. The various ecosystems within the Monument are inhabited by of protected species, including marine mam- 
mals, marine turtles and seabirds. Photos: J. Watt (right), T. Summers (center) and USFWS (left). 

Hawaiian Monk Seal 

The Hawaiian monk seal (Monachus shauinslandi) is in crisis. The population is in a decline that has lasted 
20 years, and today only about 1,200 monk seals remain. Modeling predicts that the species' population will 
fall below 1,000 animals by the year 2012. Actions to date have not been sufficient to result in a recovering 
population. Most of the population of Hawaiian monk seals breed and forage inside the Monument boundaries. 
NOAA's National Marine Fisheries Service (NMFS) is the primary federal agency responsible for the manage- 
ment of the Hawaiian monk seal and has identified the recovery of this species as the number one priority, 
based on the high magnitude of threats, the high recovery potential, and the potential for economic conflicts 
while implementing recovery actions. NMFS recently updated its Hawaiian monk seal recovery plan and has 
detailed several key actions required to address current and potential threats to the recovery and survival of 
this critically endangered species (NMFS, 2007). To advance these efforts, the Monument management board 
is pursuing several key strategies as identified in its management plan in support of monk seal recovery efforts 
(PMNM, 2008). 

Cetaceans 

Sightings and acoustic recordings of baleen whales, as well as toothed whales and dolphins have been docu- 
mented throughout the Monument. Five species of baleen whales listed as "endangered" under the ESA and 
as "depleted" under the MMPA have been sighted or heard in the Monument area. In addition to these five, the 
endangered sperm whale (Physeter macrocephalus) and at least 18 other non-ESA listed species are found 
in the Monument (see the Marine Protected Species chapter for more information). It has now been docu- 
mented that humpback whales (Megapera novaeangliae) are calving in the eastern portion of the Monument 
(Johnston et al., 2007). Recovery actions for this listed species are summarized in the final recovery plan for 
the humpback whale (NOAA Fisheries, 1991). Draft recovery plans are available for the fin whale and sperm 
whale (NOAA Fisheries, 2006a, 2006b), and a final plan is available for the recovery of the blue whale (NOAA 
Fisheries, 1998). 



Marine Turtles 

The Hawaiian green turtle (Chelonia mydas), hawksbill (Eretmochelys imbricate), loggerhead (Caretta caret- 
ta), and leatherback (Dermochelys coriacea) turtles are known to occur within the Monument boundaries. 
While there are no records of the endangered olive ridley (Lepidochelys olivacea) within Monument waters, 
their wide distribution throughout the tropical Pacific makes it plausible that they also occur there Green and 
loggerhead sea turtles are listed as threatened species; the hawksbill and leatherback turtles are classified as 
endangered species. Recovery plans and five-year reviews jointly published in 2007 are in place for each of 
these species in the Pacific (NOAA Fisheries and USFWS, 1998a; 1998b; 1998c; 1998d; 1998e, 2007). Sea 
turtle population declines have occurred across the Pacific due to nesting habitat loss, fishery interactions 
and the harvest of eggs and turtles for commercial and subsistence purposes. About 90% of the green turtles 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

in the Hawaiian Islands nest in the NWHI, the majority on a few islets at French Frigate Shoals (Balazs and 
Chaloupka, 2004). Green turtle populations have steadily increased in Hawaiian waters since the species was 
added to the list of threatened species in 1978. 

Seabirds 

Five endangered bird species in the NWHI are protected under the ESA. The only seabird occurring in the 
NWHI listed under the ESA is the Short-tailed Albatross (P. albatrus). The other four ESA listed endangered 
bird species are the Laysan Duck, Laysan Finch, Nihoa Finch and Nihoa Millerbird. 

The Short-tailed Albatross breeds primarily on Torishima, an island owned and administered by Japan. The 
Short-tailed Albatross was first observed at Midway Atoll between 1936 and 1941. Since then, one to three 
individuals have been observed every year in the NWHI, primarily on islands in the northwestern half of the 
Monument. Although short-tailed albatrosses do not currently nest in the NWHI, a small number of adult birds 
conduct breeding displays each year at Midway Atoll. The Short-tailed Albatross Draft Recovery Plan provides 
recommendations for ways in which Monument staff can facilitate recovery of this species (USFWS, 2005). 

The Laysan Albatross (P. immutabilis) and the Black-footed Albatross (P. nigripes) are both considered endan- 
gered by International Union for Conservation of Nature (IUCN) and BirdLife International. Both species breed 
in the NWHI and also forage outside of the Monument. A Conservation Action Plan for both Black-footed Alba- 
tross and Laysan Albatross (Naughton etal., 2007) has been developed to provide managers with a framework 
for the conservation of both species. In addition to the three albatross species that occur in the Monument, 
another 18 species of seabirds and five species of shorebirds that regularly breed and overwinter, respectively, 
in the NWHI, are fully protected under the MBTA. The Monument Management Plan (MMP) includes activities 
to protect and enhance seabird and shorebird habitat and to minimize impacts of habitat destruction, contami- 
nants and fisheries interactions 

Management and conservation of migratory species such as cetaceans, marine turtles and seabirds will re- 
quire coordination with international partners and conservations organizations outside of the Monument. 



MANAGEMENT OF THREATS TO THE ECOSYSTEM 

Anthropogenic activities impact oceans worldwide (Halpern et al., 2008). On a global scale, the highest rank- 
ing threats to coral reefs were identified as sedimentation, coastal development, trampling and nutrient inputs 
(Halpern et al., 2007). At a regional scale, these threats vary by location. The remoteness of the Monument as 
well as its limited emergent land results in a different suite of identified ecosystem threats. 

The Monument is affected by past changes to the ecosystem as well as current on-going threats. The emer- 
gent land areas and potentially some near shore waters continue to be affected by contaminants left over from 
military use of the islands. During World War II, large scale modifications such as channelization were made to 
the environment which changed the flow of water within the atolls and continues to impact the local ecosystem. 
In addition, marine species that also use emergent land such as seabirds are negatively impacted by invasive 
terrestrial species and other terrestrial based threats such as contaminants. It is important for mangers to be 
able to identify the potential threats and evaluate the impact of the threat to the overall functioning of the eco- 
system at a local scale as well as a regional scale. 

In a recent threat analysis of the NWHI region, 24 potential threats were analyzed based on vulnerability fac- 
tors and order of magnitude of the threat (Selkoe et al., 2008; Table 10.1). The analysis was focused on threats 
to the marine environment across the NWHI region. A systematic and quantitative method was used to collect 
and synthesize expert opinion on the ecological effects of these potential anthropogenic threats to the region. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

The following discussion of threats to the Monument focuses on the top four threats identified across ecozones 
for the region by expert opinion. These threats are: 

1. Climate change 

2. Marine debris/ghost fishing 

3. Alien species 

4. Ship groundings 



Table 10.1. Potential threats analyzed by Selkoe et a/., 2008. Source: Selkoe et a/., 2008. 



THREAT 



Alien species 
Anchor damage 



EXPLANATION 



Includes only populations that have established, not single sightings 
Includes large anchors in deepwater and small anchors of tender boats 



Aquarium collecting 



Primarily aquarium trade activities, only one example of legal event to date 



Bottomfishing 
Coastal engineering 
Diver impacts 
Ghost fishing 
Increasing UV radiation 



An ongoing fishery for a suite of deepwater snapper and grouper using hydraulic handlines >100 fath- 
oms depth outside three nautical miles. Boats also troll in transit, impacting pelagic fish and birds 

The lingering impacts of past dredging, seawalls and pier construction, and ongoing maintenance 
activities, primarily at Midway and French Frigate Shoals 

Includes diving for any purpose (but it primarily occurs for research); may cause disturbance to ani- 
mals, damage to corals, potential for inter-site transfer of micro-organisms 

A subcategory of marine debris - mostly discarded monofilament and rope nets and some lost lobster 
traps that ensnare and drown animals and smother reefs 

Increased ultraviolet radiation from the anthropogenic thinning of the ozone layer 



Indigenous fishing 



Fishing by native Hawaiians for consumption - this potential activity is likely focused on southeast end 
of NWHI 



Chemical contamination 



The leeching of chemical waste from past and ongoing military activities and habitation, primarily at 
Midway and French Frigate Shoals 



Lobster fishing 



Lobster fisheries were halted in 2000 due to population collapses. Only lingering impacts were consid- 
ered - there has been little rebound of lobster to date, potentially impacting lobster predator popula- 
tions (e.g. monk seals) 



Marine debris 



All types of man-made materials (including plastics and derelict fishing gear) that may break corals, 
entangle animals, are ingested by animals and accumulate on beaches 



Recreation 



Any recreational activities not covered by fishing and diving, such as boating, water sports, and wildlife 
viewing. 



Pelagic fishing 



Pelagic fishing is banned in NWHI waters but biological connection to the wider Pacific where long- 
lining and net fishing is intense may impact NWHI species which forage in the Pacific, both as bycatch 
and because the tuna on which some birds depend for foraging are being depleted 



Research installations Installation of equipment or otherwise modifying benthos, or disturbing animals 



Research wildlife sacrifice 
Sea level rise 

Sea temperature rise 

Sea water acidification 



Any lethal sampling of organisms for research activities 

Increased sea level from the anthropogenic warming of the planet. May alter habitat availability and 
stress populations of depth-dependent species like corals 

Increased temperature from the anthropogenic warming of the atmosphere. A suspected cause of 
increases in coral bleaching and coral disease, among other potential effects 

Decreased sea water pH due to the anthropogenic carbon loading of the atmosphere 



Ship groundings 



Includes the damage and disturbance from grounding, fuel spill, debris, cyanobacteria inoculation and 
debris removal. 



Ship pollution 



Includes the discharge of bilge water, sewage, spilled fuel, trash and noise and light pollution. Includes 
all types of vessels (fishing, research, shipping, tourist). 



Sport fishing 



Most relevant to southeast end of chain, the lingering impacts of the abated catch-and-release opera- 
tion at Midway and Pearl and Hermes, and ongoing trolling during ship transit 



Trampling damage 
Vessel strikes 



Walking on beaches, intertidal, emergent land and reef flats not associated with diving 

When a small or large vessel hits benthic communities or large mobile animals while in transit, without 
grounding 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Managers recognize the need to evaluate threats based upon the source and impact of the identified hazard. 
If a threat occurs within the management boundary, managers have more opportunity to successfully mitigate 
the threat. Threats generated outside of the management boundary still require a response, but they are often 
more difficult to prevent. Of the top four identified threats to the Monument, climate change and marine debris 
originate outside the boundaries of the Monument. Alien species establishment and ship groundings are con- 
sidered locally-based threats that occur within the Monument boundaries. 

Climate change factors are already affecting the NWHI ecosystem and will have a widespread impact. Sea- 
level rise is already impacting available habitat for species such as the Hawaiian monk seal, green turtle and 
the seabirds. The impact from sea surface temperature (SST) changes will be seen throughout the ecosystem 
including coral bleaching and potential impacts to prey species for seabirds and other predators. Marine debris 
was also identified threat from outside the Monument that will impact many different ecozones. 

Alien species and ship groundings are identified as high local threats because they have a potentially large 
impact on ecosystem function and long recovery times. To date, the threat of alien species may not be highly 
significant in marine areas within the NWHI (i.e., 13 marine alien species currently known to occur within the 
Monument) but the potential impact from an introduction could be widespread. There is always the potential for 
ship groundings but emergency response plans to minimize the impacts of the groundings are under develop- 
ment 



Selkoe et al. (2008) evaluated, eight different ecozones (Table 10.2). A finer-scale threat analysis can occur by 
examining the ecozones and assessing how a threat may impact each specific area. The inner and outer reef 
zones were identified as the most vulnerable. Emergent land was considered vulnerable because of sea level 
rise and the potential loss of habitat. The shallow water ecozones were more vulnerable to extra-boundary 
threats than local threats. Many of these top threats are difficult for local managers to control because they 
arise from activities outside the Monument boundaries, indicating that additional work is needed to preserve 
the NWHI despite its highly protected status. The analysis indicates where interagency cooperation in remov- 
ing and mitigating threats should be focused (Selkoe et al., 2008) 

Table 10.2. Ecozones evaluated in the threat analysis. Source: Selkoe et al., 2008. 



ECOZONE DESCRIPTION 



strial Interior land distinct from the littoral zone 

Rocky Inter- Solid substrate at intertidal depth composed of basalt rock 
tidal | 

Sandy Beach Intertidal beach and adjacent shallows with soft benthos 

Algal Beds Primarily Halimeda beds in lagoons and deeper terraces, but 

also small stands of endemic seagrass at Midway 

Inner Reef Refers to shallow, mostly protected reef areas (lagoonal, 

back, reticulated or patch reefs) 

Outer Reef Exposed seaward reefs from the crest down to the slope less 

than 30 m depth 

Deep Reef/ Deep reef is designated >30 m depth. Banks are sites of high 

Banks relief benthos in deep waters with rich fish communities, with 

or without reef builders 

Pelagic Waters The entire water column, from surface to depth, outside of 
lagoon and shallow reef environments 



DISTRIBUTION 



Kure, Midway, Lisianski, Lysan, Mokumanamana 
and Nihoa 

Gardner Pinnacles, La Perouse Pinnacle (French 
Frigate Shoals), Mokumanamana and Nihoa 

Kure, Midway, Pearl and Hermes, French Frigate 
Shoals, Lisianski, Laysan, and Nihoa 

Kure, Midway, Pearl and Hermes, French Frigate 
Shoals, Lisianski, Laysan, Mokumanamana and 
Nihoa 

Kure, Midway, Pearl and Hermes, Lisianski, Neva 
Shoals, Laysan, Maro, and French Frigate Shoals 

Most NWHI locations where depth is <30 m 



There are approximately 30 deep banks in the 
NWHI. Deep reef is usually found adjacent to any 
shallow reef area. 

Makes up the majority of NWHI habitat. 



Climate Change 

Sea level rise, changing storm intensity and frequency, sea surface temperature (SST) rise and acidification 
are components of climate change most likely to affect the Monument. Evidence of sea level rise has already 
begun to adversely affect the available terrestrial habitat and models predict that sea level will continue to rise. 
SST is monitored via satellite in addition to using buoys at several locations throughout the NWHI, resulting 
in a long-term temperature time series for Midway Atoll (Jokiel and Brown, 2004). Elevated SST has already 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

impacted corals as indicated by recent coral bleaching events in the Monument (see Coral Bleaching for more 
details). With regards to ocean acidification, the third component of climate change most likely to affect the 
Monument, staff members are in the process of designing experiments to characterize the carbonate chemis- 
try and establish a baseline for the NWHI. 

Sea Level Rise 

Global mean sea levels have risen an estimated 3.1 ± 0.7 mm yr _1 from 1993-2003, an amount higher than any 
other 10-year period since 1950 (IPCC, 2007). However sea level rise varies regionally and in order to under- 
stand the effects on the ecosystem it must be monitored at the island and atoll scale. 

One effect of rising sea level in the NWHI is the loss of habitat. Emergent land in the NWHI is estimated at a 
total 14 km 2 and the loss of available emergent land will greatly reduce the available habitat for many species. 
The effect of habitat loss on species that use emergent land features will impact many of the species that are 
already rare and maybe devastating to those populations that depend on these islands for survival. Marine 
species that will be impacted by sea level rise include the Hawaiian monk seal, green turtle and several sea- 
bird species. In addition there is the potential for further habitat degradation with the release of contaminants 
contained in landfills and other areas as the islands are eroded or flooded from sea level rise. 



SCENARIO 


RISE LEVEL 


BASE SEA LEVEL 


Scenario 1 - Low 


9 cm 


Mean Low Water (MLW) 


Scenario 2 - Low 


Spring Tide 


Scenario 3 - Median 


48 cm 


Mean Low Water (MLW) 


Scenario 4 - Median 


Spring Tide 


Scenario 5 - High 


88 cm 


Mean Low Water (MLW) 


Scenario 6 - High 


Spring Tide 



Evidence of sea level rise can be clear- Table 10.3. Sea level rise scenarios modeled for French Frigate Shoals, 
ly observed with the submersion of Lisianski Island and Pearl and Hermes Atoll. Source: Baker etal., 2006. 

Whaleskate Island within French Frig- 
ate Shoals in the late 1990s (Baker et 
al., 2006). NOAA Pacific Islands Fish- 
eries Science Center (PIFSC) modeled 
the potential terrestrial habitat loss from 
sea level rise using estimated sea level 
rise values and current elevation data 
collected in the field. The study included 
islands within Pearl and Hermes Atoll, Lisianski Island and French Frigate Shoals. Pearl and Hermes Atoll 
and French Frigate Shoals are both composed of small low lying islets surrounded by a barrier reef, whereas 
Lisianski Island is a single large, low-lying island (Figure 10.5). Six different scenarios using three sea level 
rise values and two different tide conditions were evaluated (Table 10.3). The results of the modeling indicate 
that sea level rise will affect each island group differently. Lisianski Island (the largest and highest island in the 
analysis), could experience the least amount of impact with only a 5% decrease in area using the highest sea 
level rise scenario. In contrast, the highest seal level rise scenario for the islets at Pearl and Hermes Atoll and 
French Frigate shoals range from a 25% loss up to 90% loss (Baker et al., 2006). The estimates produced by 
the model were based on the assumption that the island shape remains constant and the model did not include 
erosion factors. Efforts to develop a monitoring system of the changes in the size of the islands are currently 
underway. 

Sea turtles are dependent on terrestrial areas for nesting. Islets in French Frigate Shoals support the majority 
of the Hawaiian green turtle breeding population. The loss of habitat and the possibility for nesting areas to 
be flooded during nesting times could have a large impact on the species. In other regions, green turtles have 
demonstrated intra-specific nest destruction once habitat is lost and nest density becomes high (Bustard and 
Tognetti, 1969). This behavior may occur among nesting Hawaiian green sea turtles in the Monument if nesting 
habitat is destroyed. 



Impacts to seabirds will vary depending upon the species. Over 90% of Black-footed and Laysan Albatrosses 
breed in the Monument and the loss of habitat in this area could affect the overall world population. The largest 
breeding populations of these albatross species occur at Midway Atoll and Laysan Island which were not ana- 
lyzed in the sea level rise study, but habitat loss is expected to be similar to Lisianski Island because they are 
both large islands. The predicted decrease in area would not be large, but the species remain vulnerable as 
the islands represent such a critically important breeding area. Other seabird species may also be potentially 
impacted by the loss of nesting habitat as the result of climate change. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



510 




Lisianski Island 



B 



East Island (FFS) 




O Interpolated points 
A Theodolite stations 
A Mean low water points 



Legend 




Median rise + spring tide 
0.89 (MLW + maximum rise) 
0.49 (MLW + median rise) 
0.10 (MLW + minimum rise) 
MLW 



120 



urn 




Trig Island (FFS) 



D 




740 




llll 




m 


fh 




•-." 


w ^^ 


(n) 


Southeast Island (PHR) 



Figure 10.5. Current and projected maps of four NWHI at mean low water (MLW) with minimum (9 cm), median (48 cm) 
and maximum (88 cm) predicted sea level rise. The median scenario at spring tide is also shown. (A) Lisianski Island; (B) 
East Island, French Frigate Shoals, showing the measured and interpolated points along the waterline and berm used 
to create the Triangular Irregular Network (TIN); (C) Trig Island, French Frigate Shoals; (D) Southeast Island, Pearl and 
Hermes Reef. Source: Baker et al., 2006. 




W 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Storm Intensity and Frequency 

Global weather patterns appear to be changing and climatologists suggest that the increasing intensity and 
frequency of storms may be related to modern, anthropogenic influences (IPCC, 2001; Nott, 2003; Nott, 2004; 
Nott et al., 2007). Studies into storm trends have shown that while SSTs have been rising, the intensity of the 
storms occurring in this time period has increased (Hoegh-Guldberg, 1999; IPCC, 2001; Hughes, 2003; Web- 
ster et al., 2005; IPCC, 2007; NOAA Satellite and Information Service, 2007). While sheer numbers of tropical 
storms have remained relatively constant, the destructiveness of the cyclones has increased over the past 30 
years and hurricanes in the Category 4 and 5 range have nearly doubled since the 1970s (Emanuel, 2005; 
Webster et al., 2005). If climate patterns follow the projections, storms may continue to increase in intensity 
over the coming years. As storm intensity increases, the impacts felt by low-lying islands and atolls could prove 
detrimental to the inhabitants. 





Mm 



The damage from high intensity storms 
to low lying sand islands was demon- 
strated in late December 2008 at French 
Frigate Shoals when high levels of ero- 
sion occurred at East Island. Using re- 
mote cameras available for monitoring 
turtle nesting activity scientists at PIF- 
SC were able to evaluate the damage to 
East Island which included several feet 
of erosion along the northwest side of 
the island (G. Balazs pers. comm; Fig 
10.6). Monitoring will need to be con- 
ducted to evaluate the long-term impact 
of the erosion event and to determine if 
accretion will occur to other parts of the 
island. Researchers active in the NWHI 
over the past 30 years have observed 
changes to the size and shape of many 
of the islands (G. Balazs and J. Mara- 

gos, pers. comm.). In order to determine if these observed changes are from sea level rise or storm damage a 
monitoring program will need to be developed. 




:"•■ .' 

-I --• • \'t .* • 

: > w •■: .j ,- ... 






Figure 10.6. High levels of erosion on East Island just days after highly 
intense storms passed through the area in December 2008. Photo: PIFSC 
and G. Balazs. 



Sea Surface Temperature Change 

Eleven of the years spanning 1995 to 2006 are ranked among the warmest 12 years of recorded global sur- 
face temperature (IPCC, 2007). Temperature change is another component that may impact the Monument's 
marine ecosystems. The NWHI are monitored as part of the Coral Reef Early Warning System (CREWS). The 
system provides managers and researchers with telemetered meteorological and oceanographic data at pre- 
cise locations. In the NWHI, NOAA's PIFSC, Coral Reef Ecosystem Division (CRED) has deployed long-term 
moored observing stations, satellite-tracked drifting buoys, and subsurface instrumented moorings (Table 10.4 
and Figure 10.7) Changes in SSTs will result in changes to available habitat for temperature dependent spe- 
cies and coral bleaching. 

As SSTs change species currently using habitat near the surface may move to lower depths or different lati- 
tudes to find the appropriate habitat conditions. This will end up impacting other species dependent on these 
species. For example, seabirds feed on fish and other marine species near the ocean surface. As SST in- 
creases, seabird prey species move to deeper, cooler water, decreasing food availability for foraging birds, or 
requiring birds to fly further north in the Pacific to obtain food resources. 



Changes in SST poses a threat to coral reef ecosystems in the form of coral bleaching. Corals are symbiotic 
organisms which secrete a hard, mineral calcium carbonate structure. The symbiosis is between microscopic, 
photosynthetic organisms called zooxanthellae that inhabit the soft tissue of the coral polyp. Zooxanthellae 
provide the pigmentation of the coral and produce energy which is donated to the host and contributes sig- 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Table 10.4. Distribution of long-term oceanography monitoring buoys in the Monument. 





CREWS-ENH 1 CREWS-STD 2 


SST-ARGOS 3 


ODP 4 


WTR 5 




Kure 


-- 


X 


X 


-- 


X 


X 


Midway 


-- 


-- 


X 


X 


-- 


X 


Pearl and Hermes 


-- 


X 


-- 


X 


X 


-- 


Lisianski 


-- 


-- 


X 


-- 


X 


X 


Laysan 


-- 


-- 


X 


-- 


-- 


X 


Maro Reef 


-- 


X 


-- 


-- 


-- 


X 


Gardner Pinnacles 


-- 


-- 


-- 


-- 


-- 


X 


French Frigate Shoals 


-- 


-- 


-- 


-- 


-- 


X 


Mokumanamana 


-- 


-- 


-- 


-- 


-- 


-- 


Nihoa 


-- 


-- 


-- 


-- 


-- 


-- 



^oral Reef Early Warning System Enhanced - Moored buoys which provide high resolution SST, barometric pressure, wind speed, 
wind direction, and additionally provide salinity, UV-B, and PAR. 

2 Coral Reef Early Warning System Standard - Moored buoys which provide high resolution SST, barometric pressure, wind speed 
and wind direction. Subsets of these data are transmitted daily via satellite telemetry. 

3 Moored buoys which provide high resolution SST. Subsets of these data are transmitted daily via satellite telemetry. 
"Subsurface Ocean Data Platform - Subsurface moorings, providing high resolution current profiles, directional wave spectra, and 
temperature and salinity. 

5 Subsurface moorings providing high resolution wave and tide records, temperature and conductivity. 

6 Subsurface moorings providing high resolution temperature. Additionally used on towed platforms for temperature and pressure- 
based depth. 



nificantly to the ability of a coral to grow 
and reproduce. When a coral is stressed 
by higher than normal temperatures, 
sometimes as little as a 2-3°C increase 
in temperature above their optimal tem- 
perature, they expel their zooxanthellae 
into the water column resulting in a loss 
of color (Hoegh-Guldberg, 1999). In this 
bleached, energy depleted state a coral 
is more susceptible to disease infiltra- 
tion and overgrowth by fast-growing turf 
algae. Anthropogenic activities resulting 
in increased nutrient loads, sedimenta- 
tion and physical damage at the site can 
make bleaching events worse. 

SST anomalies resulting from regional 
and global-scale climatic phenomena 
are believed to be the cause of bleach- 
ing in the NWHI. Mass coral bleaching in 
the NWHI occurred during late summer 
2002 (Aeby et al., 2003; Kenyon and 

Brainard, 2006). This was the first ever recorded bleaching event known to occur in the NWHI. Coral bleach- 
ing occurred again at high levels in 2004, and was detected again at low rates in 2006 (Kenyon and Brainard, 
2006). The corals in the NWHI were believed to be less susceptible to bleaching due to the high latitude loca- 
tion. Bleaching was most severe, however at the three northernmost atolls (Pearl and Hermes Atoll, Midway 
Atoll and Kure Atoll), which experience both higher and lower SSTs than other reefs of the NWHI. During the 
bleaching event, greater magnitude and longer durations of temperature anomalies were recorded and attrib- 
uted to the bleaching events of 2002 (Hoeke et al., 2006). Lisisanski, Laysan and Maro experienced shorter 
and less severe temperature anomalies, resulting in comparatively minor bleaching events. Field investiga- 
tions conducted in 2004 indicate that bleaching occurrence was highest in shallow back reef and patch reef 
habitats (Kenyon and Brainard, 2006). Researchers from the University of California, Santa Cruz are currently 
modeling circulation patterns at Midway Atoll to determine if anthropogenic changes to the flow and circulation 
patters may exacerbate bleaching events. 




Legend 

■ Oceanography Data Collection Si'es 
; I Papahanaumokuakea M 

A 



Marine National Mtonumenl 

100 200 300 400 

u Kilometers 



Figure 10.7. Locations of oceanography monitoring buoys in the NWHI. 
Map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



In addition to using field measure- 
ments it is possible to detect potential 
coral bleaching events using satellite 
information. NOAA's Coral Reef Watch 
produces near-real-time alerts from 24 
selected reefs around the globe. Mid- 
way Atoll is one of the sites that is moni- 
tored. Figure 10.8 contains graphs indi- 
cating when Midway Atoll SSTs reached 
levels that are associated with beaching 
events. The graphs show the 2002 and 
2005 SST associated with bleaching 
events were detected by NOAA's Coral 
Reef Watch prior to being documented 
in the field. 

The development of a bleaching re- 
sponse plan is critical to effective reef 
management. This ensures that when 
a bleaching event occurs, decisive ac- 
tion can be taken as soon as possible 
to mitigate the effects of the bleaching 
event. The NWHI will require a unique 
response plan given that the region is 
largely free from local sources of an- 
thropogenic stressors, so bleaching 
is largely a result of increased SSTs. 
However, managers will work to iden- 
tify resilient areas based upon the best 
available information from monitoring 
data, research projects, past bleach- 
ing events and modelling. A bleaching 
response plan would call for identifica- 
tion of specific groups and actions that 
should take place prior to a bleaching 
event, during the event and follow up 
after SSTs have returned to normal. 

Ocean Acidification 

Coral reef systems maintain a deli- 
cate balance between calcification and 
erosional forces. For the reef to grow 
and accrete mass, the corals' ability to 
calcify must outrun the pressures put 
upon the system, such as bioerosion, 
physical erosion from wave action and 
storms and anthropogenic damage. 
The NWHI are relatively shielded from 
most anthropogenic effects due to their 
remoteness, but they are still subject to 
natural forces. The shallow and deep 
water coral reefs in the NWHI will po- 
tentially be impacted by changes in the 
carbonate levels in the ocean. If the reef 



25 



T "--- 15 : 









Midway Atoll, US (Jan 


1 2000 - Dec 31 2001) 








j a 




- Bleaching Threshold SST Max Monthly Mean SST + Monthly Mean Climatology 4, B DHW3 _ 


30 
























25 








+ • + \_ 










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20 


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+ 




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+ '■ 








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J FMAMJJASONDJ FMAMJJASOND 
2D0D 2D01 





□ No Stress □ B 


eaching Watch □ Bleach 


ing Warning ■ Alert Level 1 


□ Alert 


Level 2 




Midway Atoll, US (Jan 1 2002 - Dec 31 2003) 


JS 


~ Bleaching Threshold SST Max Monthly Mean SST + Monthly Mean Climatology 4, B DHWs ~ 


30 


















: 




-- - -: 





rP 1 - ^"^i. ~ ^ ~ '■ 


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25 


















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20 
15 




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-" 


--!-- 


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2D02 2D03 

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Midway Atoll, US {, 


an 


1 


2004 - I 


ec 


31 


2005) 






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— Bleaohing Threshold SST Max Monthly Mean SST + Monthly Mean Climatology 


4, B DHWs : 


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: 






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FMAMJJASONDJ FMAMJJASON 

2D04 2DCI5 

□ No Stress □ Bleaching Watch □ Bleaching Warning ■ Alert Level 1 ■ Alert Level 2. 







vlidwaj 


Atoll, US (Jan 


1 


2006 - Dec 


31 2007) 




M 


~ Bleaohing Threshold SST Mdx Monthly Mean SST + Monthly Mean Climatology - 


- 4 B DHWs I 


30 


- i i 














I 1 




- _:_ _; 





|C/Vf —i 




4. 


j^r^ 


i^_ ; _ 


25 


- i : 










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+\-^ 


20 


- ■ +: + 


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._.. 


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ii i ii i 



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: 20 

D 

x 

15* 

o 

10 S 
■s 
- 



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: 20 
15 

re 
- 



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J FMAMJJASONDJ FMAMJJASOND 
2006 2007 







□ No Stress 


22 Bleaching 


Watch □ B 


ea cl- 


ing 


*arn 


"ng 


□ Alert Level 1 


5 Alert 


Leve 


2 








Midway Atoll, US (Jan 1 200S - Nov 6 2008) 


JS 




- Bloaohing Threshold SST Man Monthly Mean SST 4 Monthly Mean Climatology 4, B DHWs I 


30 




































i 







h 




_ ^ _ 




„L^_ 








_ 










l_ 









25 


























+ 






+ 




i 


20 


I_ 


+ 
+ + 




;.t; 


;; 


- 






+ 


+ 


+ 


-; - 


;_ 


;_ 


;;: 


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: 


_+- 


15 






































i MM 



25 

20 

□ 

x 

15* 

o 
10 S 



- 5 



J FMAMJJASONDJ FMAMJJASOND 

20DS 20D9 

□ No Stress □ Bleaching Watch □ Bleaching Warning S Alert Level 1 H Alert Level 2 



Figure 10.8. Sea surface temperature values recorded at Midway Atoll 
documenting potential conditions for coral bleaching events (2000 - 2008). 
Source: NOAA Coral Reef Watch. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

structure changes there can be will be an ecosystem wide effect as habitat availability and species ranges 
change. 

Evidence of undersaturation has been reported in the Intermediate Pacific in relatively shallow waters of be- 
tween 200 m and 1,000 m (Feely et al., 2002). This is of concern to the NWHI deepwater habitat which for 
these purposes will be defined as depths greater than 50 m. A number of corals have identified habitat ranging 
from 20-3,000 m (Hourigan et al., 2008). These corals provide structure to the benthos as well as serving as 
habitat for other organisms living in the deep that have yet to be fully explored. Endangered Hawaiian monk 
seals with attached animal borne imaging systems (Crittercams) have been recorded foraging for fish that find 
shelter in black coral beds (Parrish and Baco, 2008). However, due to the remoteness of the NWHI follow-up 
deepwater surveys have yet to be conducted, as they require extensive time, planning and budgeting con- 
siderations. Deepwater corals are mainly slow-growing species, and the effects of undersaturation may not 
manifest for years after it has occurred. Or, conversely, if the saturation horizon rises to shallow depths, the 
aragonite structure of the deepwater corals may begin to dissolve away into the ocean. 



The degree to which ocean acidification will affect Monument coral reefs is presently under investigation. Plans 
are underway to establish a baseline of the carbonate chemistry of the NWHI. This would be done by utiliz- 
ing the CTD (conductivity, temperature, depth) sensor scanner onboard R/V Hiialakai and outfitting it with a 
pH sensor. Carbonate chemistry of a water sample can be characterized by taking three measurements; the 
dissolved inorganic carbon, alkalinity and pH. All three of these measurements are planned to be performed 
onboard during regular CTD casts and the seawater samples brought back for further analysis. These types 
of measurements, taken from deepwater habitats and lagoon waters (from small boat platforms) will help the 
Monument to develop an understanding of the current carbonate chemistry of the waters and allow us to moni- 
tor future changes. 



Ocean Acidification 

Ocean acidification is the process of seawater becoming less basic and is likely to upset the delicate balance between 
reef calcification and erosion (Hoegh-Guldberg, 1999). The carbonate equilibrium describes the process of calcification in 
marine animals and is illustrated by equation 1 (Kleypas et al., 1999; Royal Society, 2005; Kleypas and Langdon, 2006). 
Acidification, where atmospheric C0 2 is absorbed by the surface of the ocean where it forms carbonic acid, is illustrated 
by equation 2;. This acid then dissociates into free hydrogen and bicarbonate ions resulting in increasing amounts of bi- 
carbonate, leaving less carbonate ions available to interact with the abundant calcium ions present in seawater. Calcium 
ions are not thought to be a limiting factor in calcification (Royal Society, 2005). Therefore, when the amount of C0 2 in the 
atmosphere increases, the availability of carbonate ions for calcifying organisms (corals, calcareous algae, plankton etc.) 
to incorporate into their skeletons decreases and the carbonate equilibrium shifts facilitating the dissolution of the calcium 
carbonate skeleton (Kleypas et al., 1999; Royal Society, 2005; Kleypas et al., 2006). 

Equation 1: The Carbonate Equilibrium 

<- Skeletal formation 



Ca 2+ + 2HCO, 



CaC0 3 + C0 2 + H 2 

Skeletal dissolution 



Equation 2: Acidification 



C0 2 + H 2 



H 2 CQ 3 



H + +HCO, 



H + + C0 3 2 " 



There are two forms of calcium carbonate found in skeletons of CaC0 3 secreting organisms. The first is calcite, which is 
the less soluble form found in crustose coraline algaes, and the other is the aragonite form of calcium carbonate, which is 
utilized by scleractinian corals and other pteropods. There is a critical concentration of carbonate ions in seawater below 
which calcium carbonate will dissolve. The solubility of calcium carbonate is a function of depth and pressure. The critical 
concentration occurs at a depth known as the "saturation horizon", under which calcium carbonate structures tend to dis- 
solve. Due to increased amounts of C0 2 in seawater and the resultant decrease in the carbonate ion concentration (equa- 
tion 2) the saturation horizon will move ever shallower with increasing releases of anthropogenic C0 2 into the atmosphere. 
This means that the depth at which corals are able to calcify will grow shallower as more C0 2 is input into the atmosphere. 
Latitude also plays a role when looking at where saturation boundaries occur. Lower latitudes near the equator tend to 
have saturation states conducive to the solidified structure of corals, while poleward areas are already showing evidence 
of undersaturation at the surface waters. Currently the surface waters of the NWHI still fall within saturation parameters 
(Feely etal., 2002). 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Marine Debris 

A multiagency effort initially launched in 
1996 by the University of Hawaii's Sea 
Grant College Program began to ad- 
dress the problem of marine debris, a 
problem that was much larger than any 
one agency alone can resolve. An esti- 
mated 750 to 1,000 tons of marine de- 
bris were on reefs and beaches in the 
NWHI (NOAAPIFSC, unpublished; Fig- 
ure 10.9). NOAA, in collaboration with 
14 other partners including the U.S. 
Coast Guard (USCG), Schnitzer Steel 
Hawaii Corporation (formerly Hawaii 
Metals Recycling Company), the Hawaii 
Sea Grant College Program, U.S. Navy, 
USFWS, the City and County of Ho- 
nolulu, the state of Hawaii, The Ocean 
Conservancy, Hawaii Wildlife Fund, 
Matson Navigation Company, and oth- 
ers removed 66 tons of marine debris 
and derelict fishing gear from 1996 to 
2000. In 2001, the multiagency cleanup 
effort was extended, resulting in a cor- 
responding increase of marine debris 
removed from reefs and beaches of the 
NWHI (Table 10.5). The total amount 
of marine debris removed from 1996 to 
2007 was 582 tons. 

The source of much of the marine de- 
bris is fishing nets discarded or lost in 
the northeastern Pacific, well outside 
of the Monument boundaries. In order 
to address the source of marine debris 
in the Pacific, Monument managers will 
need to work with international partners 
to look at methods and develop policies 
for reducing marine debris. Even if all 
new input of debris were stopped, exist- 
ing debris in the ocean would continue 
to accumulate in the NWHI for years to 
come. At a Pacific basin scale, it is sug- 
gested that the subtropic convergence 
zone (STCZ) that moves between 25°N 
and 35°N is an area of high ghostnet re- 
tention (Figure 10.10). When the STCZ 
moves within range of the NWHI, the 
nets often become entangled on reefs 
and continue to be an entanglement 
hazard for many species. Once the de- 
bris reaches the NWHI, the rate of ac- 
cumulation of nets on reefs varies by is- 
land and atoll and within island and atoll 




Figure 10.9. Divers cutting away nets from the reef in the NWHI. Photo: 
CRED. 



Table 10.5. The total amount of marine debris removed from 1996 to 2007 
was 582 tons. Source: NOAA/PIFSC. 


YEAR TONS REMOVED 




1996 - 2000 


Approximately 25 tons per year 


2001 


68 


2002 


107 


2003 


118 


2004 


126 


2005 


57 


2006 


21 


2007 


59 




Figure 10.10. Map illustrating regional current movements and the conver- 
gence zone. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

(Dameron et al., 2007). A model of potential accumulation rates was developed for the NWHI to help evaluate 
the distribution of marine debris and where efforts should be focused for removal. The results from the model 
can be used by managers to target areas for marine debris clean-up efforts. 



Table 10.6. 2007 field season marine debris removal by island and atoll. 
Source: 



In the 2007 field season Kure Atoll, 
Pearl and Hermes Atoll, Lisianski Is- 
land, Laysan Island and French Frigate 
Shoals were targeted for marine debris 
removal (Table 10.6). In addition to the 
marine debris removal efforts, a study 
of the effects of marine debris and ma- 
rine debris removal on NWHI coral reef 
benthic communities was initiated at 
Midway Atoll in 2008. The study will be 
assessing the long-term effects of re- 
moval of nets from reefs as well as the 
effects of nets left in place. Initial survey 
and removal efforts began in August 
2008 and the study will continue into 
2009. The results of the study will help 
managers develop better guidelines for 
marine debris removal and decreasing 
overall impact to the coral reef ecosys- 
tems. In addition the results will begin to provide managers with information about recovery rates of the benthic 
communities following debris removal or other anthropogenic disturbance. 



LOCATION 




DEBRIS TYPE 


REMOVED 

(kg) 


REMOVED 
(tons) 


French Frigate Shoals 


Marine Debris 


5,554 


6 


Land Debris 


1,735 


2 


Kure 


Marine Debris 


2,860 


3 


Land Debris 


1,431 


1 


Laysan 


Marine Debris 








Land Debris 


2,073 


2 


Lisianski 


Marine Debris 








Land Debris 


4,396 


4 


Pearl and Hermes 


Marine Debris 


39,250 


39 


Land Debris 


1,911 


2 


Total Marine Debris Weight 


47,664 


48 


Total Land Debris Weight 


11,546 


12 


Total Debris Weight 


59,210 


59 



Marine Alien Species 

Monument managers have identified marine invasive species including pathogens as a significant threat and 
are taking action to prevent any additional introductions. Because of the Monument's vast size, it is difficult 
to carry out surveys to detect marine invasive species. However, based on the few surveys conducted (see 
Nonindigenous and Invasive Species Chapter), there are currently 13 marine invasive species that have been 
identified and documented in the NWHI. Compared to the 343 marine invasive species that have been identi- 
fied and documented in the Main Hawaiian Islands (MHI), the NWHI have a relatively low abundance of in- 
vasive species (Eldredge and Carlton, 2002; Godwin et al., 2006; Godwin, 2008). The potential of additional 
introductions of non-indigenous species in the NWHI could have dramatic consequences to the ecosystem. 
Management tools to reduce the potential introduction and spread of alien species in the Monument are the 
permitting process, enforcement of regulations and development of a monitoring and research program. 

Mandatory hull inspections for all permitted vessels are the primary tool managers can use to reduce the po- 
tential of marine alien species introductions. Prior to receiving a Monument permit, any ship that has applied 
for a permit to access the Monument must complete a hull inspection. Another mechanism for potential intro- 
ductions is byway of ballast water exchange. In response to national concern regarding invasive species, the 
National Invasive Species Act of 1996 was enacted. The Act reauthorized and amended the Nonindigenous 
Aquatic Nuisance Prevention and Control Act of 1990. In addition to the Monument discharge regulations, bal- 
last water exchange in the Monument is regulated by the USCG which codified a national mandatory ballast 
water management program for all vessels equipped with ballast water tanks that enter or operate within U.S. 
waters. These regulations also require vessels to maintain a ballast water management plan that is specific for 
that vessel and that assigns responsibility to the master or an appropriate official to understand and execute 
the ballast water management strategy for that vessel. 



There have been several reports written recently on the topic of marine invasive species in the NWH I (Eldredge, 
2005; Godwin et al., 2006; and Godwin, 2008). They provide a number of recommendations for managers to 
deal with invasive species. These reports provided the foundation for the prohibitions on ballast discharge in 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

the Proclamation and the actions outlined in the MMP Alien SpeciesAction Plan. The Alien Species Action Plan 
addresses prevention, monitoring of alien species, and education of Monument users and the public about the 
need to prevent alien species introductions. The following section focuses on how managers can utilize the 
geographic locations of known marine invasive species to shape their actions in three distinct realms: monitor- 
ing, permitting and research. 

Monitoring 

The locations of existing alien species populations can provide a foundation for future monitoring efforts. Moni- 
toring of existing infestations and identification of new infestations is a key component to the Alien Species 
Action plan in the MMP. In the past, relatively few marine invasive surveys were conducted and usually only 
once and often opportunistically. The MMP calls for the need to establish a coordinated and systematic effort 
to survey distributions and populations of known alien species at regular intervals. Understanding where exist- 
ing populations are can help guide managers in the development of monitoring protocols for the detection and 
potential eradication of current populations as well as future infestations. The monitoring program will provide 
important information on the spatial distribution, spread and population sizes of marine alien species within the 
Monument. Currently the existing Rapid Assessment and Monitoring Program monitoring does not specifically 
target alien species, but alien species would be identified during the MMP proposed alien species monitoring 
surveys. 

Permitting 

Knowing the locations of marine invasive species can assist managers in making decisions regarding the is- 
suance of permits in the Monument. An understanding of potential invasive vectors, combined with the knowl- 
edge of where marine invasive species reside in the MHI and NWHI will allow managers to take steps to 
minimize those vectors. Steps to minimize the introduction of potential invasive vectors can most effectively be 
implemented through the permitting process. 

Activities authorized under a Monument permit can be structured so that vessels visit the islands in an order 
that minimizes the risk of transporting invasive species. The more pristine sites should be visited first, and the 
most invaded locations last, to minimize the likelihood of organisms being transported from invaded sites to 
pristine areas. This may not always be possible but can be implemented where possible. These recommenda- 
tions are currently being done on an informal basis through consultations with the applicant, but they could be 
formally incorporated as permit conditions in the final permit. 

Managers can also restrict where vessels anchor or identify a route through the NWHI so as to minimize the 
risk of spreading invasive species through hull fouling or ballast water. To make effective decisions that can be 
justified, managers must have good geographic information on the locations of marine invasive species within 
the Monument. 

Research 

Information about where marine invasive species reside in the Monument is needed to inform managers and 
can serve as a guide on how best to direct management-driven research. Research about the effects of inva- 
sive species can be effectively targeted to the locations where known populations of those species do or do not 
exist. Accurate information about the abundance and positions of invasive species can help prioritize research 
based on species or sites. Research should be directed at the more abundant species, or the species in more 
vulnerable sites. In addition, other research that should be undertaken includes factors that cause alien spe- 
cies to become invasive and the interactions between native and alien species. 

Vessel Hazards 

With the exception of a few small boats at Midway Atoll, French Frigate Shoals, Pearl and Hermes and Kure 
Atoll, no vessels have home ports in the NWHI. Therefore, almost all marine traffic in the waters surrounding 
the NWHI is the result of transiting merchant vessels, research ships, fishing vessels, cruise ships, USCG 
ships and recreational vessels which visit infrequently. An estimated 50 vessels pass through the U.S. EEZ 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

surrounding the NWHI each day (Franklin, 2008). Vessels entering shallow waters intentionally or unintention- 
ally have a higher risk of impacting resources. 



Hazards to shipping and other forms of maritime traffic such as shallow submerged reefs and shoals are 
inherent in the NWHI's 1,931 km of islands and atolls. The region is exposed to open ocean weather and 
sea conditions year-round, punctuated by winter severe storm and wave events. Vessel groundings and the 
release of fuel, cargo and other items pose real threats to the NWHI. A number of factors have contributed to 
vessel groundings and cargo loss over 
the years. These factors include human 
error, lack of appropriate navigational 
practices, inaccurate nautical charts, 
and treacherous conditions due to low- 
lying islands, atolls, and shallow pinna- 
cles and banks. Periodically, accidental 
loss of cargo overboard causes marine 
debris or hazardous materials to enter 
sensitive shallow-water ecosystems. 

The history of shipwrecks and ground- 
ings is as old as the history of ships in 
the NWHI. Many islands and atolls are 
named for ships that went aground. In 
the last 50 years this history has con- 
tinued, with several vessel groundings 
(Figure 10.11). Most recently the Para- 
dise Queen and Grendel went aground 
at Kure Atoll in 1998 and 2007, respec- 
tively, and the Swordman II and Casitas 
went aground at Pearl and Hermes Atoll 
in 2000 and 2005, respectively. 




Legend 

I I Papaharsaumokuakea 

Marine National Monument 



400 

]K lometers 



Figure 10.11. Groundings in the NWHI in the last 60 years. Map: K. Keller. 



Unexploded ordnance, debris and modern shipwrecks, such as the fishing vessels Houei Maru #5, the Para- 
dise Queen II at Kure Atoll or the tanker Mission San Miguel lost at Maro Reef, are not protected as maritime 
heritage resources and represent a more immediate concern as threats to reef ecosystems. Mechanical dam- 
age from the initial grounding, subsequent redeposition of wreck material by storm surge, fishing gear dam- 
age to reef and reef-associated organisms, and release of fuel or hazardous substances are all issues to be 
considered in protecting the integrity of the environment. Dissolved iron serves as a limiting nutrient in many 
tropical marine areas and tends to fuel cyanobacteria (blue-green algae) or other iron limited species growth 
when the iron begins to dissolve and corrode. This is a problem particularly on atolls and low coral islands 
where basaltic or volcanic rock is absent in the photic zone and natural sources of dissolved iron in seawater 
are minimal. Therefore, any ships left in place would be an iron source that could contribute to potential cy- 
anobacterial blooms. It has been demonstrated that not removing nonhistoric steel vessels will have long-term 
detrimental effects, which in most cases can be worse than any short-term damage to the environment caused 
by the removal action. Vessel traffic can also affect natural resources through direct damage to the reef from 
anchors, waste discharge, light and noise. Monument regulations which prohibit anchoring on or having a ves- 
sel anchored on any living or dead coral to prevent anchor damage to reefs. Discharge of waste in the Monu- 
ment is also regulated by Proclamation and permit requirements. 

The designation of the PSSA and expansion of the ATBA is intended to reduce the potential for large vessel 
groundings within the Monument. In near shore areas, the mandatory requirement for a vessel monitoring sys- 
tem will allow better tracking of permitted vessels as well as provide information for emergency response thus 
reducing any potential impact from vessel grounding. Reducing the response time when groundings do occur 
will also minimize environmental impacts. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

MANAGEMENT OF HUMAN IMPACTS 

There is a long history of managing impacts from human activities in the NWHI. Beginning in the early 1900s, 
several federal and state agencies, including the Department of Defense, Department of Agriculture, Depart- 
ment of the Interior, the state of Hawaii, and the Department of Commerce were assigned protective responsi- 
bilities in the NWHI. Additionally, military defense needs during and after World War II required the construction 
of facilities and the presence of U.S. Navy and U.S. Air Force, and USCG stations on several islands in the 
northwestern archipelago through the end of the 20th century. The following figures (Figures 10.12 and 10.13) 
indicate periods of protective responsibilities of the various federal and state agencies and the time periods in 
which military presence occurred in the NWHI. The following section focuses on the management of human 
activities since the designation of the Monument. 



East Island, French Frigate Shoals temporary base 
camps established for Naval ship and aircraft 
exercises 



Midway Naval Air Station established for protection 
of U.S. during World War II 



East Island, French Frigate Shoals Coast Guard 
Long Range Navigation (LORAN) Station built 



Tern Island, French Frigate Shoals Coast Guard 
LORAN Station built 



Tern Island, French Frigate Shoals Pacific Missile 
Range Facility established 



Midway Naval Air Station downgraded to Naval Air 
Facility 



Midway Naval Air Facility decommissioned 



is !2- 

li 10 



19 6- Southeast Island, Pearl and Hermes Atoll temporary 
19§7 base camps established for Naval ship and aircraft 

exercises 



li 10 



IS \2 



K. 



li 13 



IS [6 



Green Island, Kure Atoll Coast Guard LORAN Station 
decommissioned 



l< 52 



is so 



is il 



19 3 



IS 78 



IS '9 



IS 53 



Green Island, Kure Atoll Coast Guard LORAN 

Station built 



Tern Island, French Frigate Shoals Coast Guard 
LORAN Station decommissioned 



U.S. Navy defeated Imperial Japanese Navy during the 

Battle of Midway 



Tern Island, French Frigate Shoals Naval Air Station 

built 



Tern Island, French Frigate Shoals Naval Air Station 

decommissioned 



Tern Island, French Frigate Shoals Pacific Missile 
Range Facility decommissioned 



Figure 10.12. History of military presence in the NWHI. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Midway Atoll secured as a U.S. possession by 
President Theodore Roosevelt. 

The U.S. Navy was assigned stewardship 
responsibilities for the wildlife and habitat of Midway 
Atoll. 



1! )3 



1£ 19 



Hawaiian Islands National Wildlife Refuge established 
by President Franklin D. Roosevelt. 

U.S. Fish and Wildlife Service was assigned 
management responsibilities for the previously- 
designated Hawaiian Islands Bird Reservation as its 
status was changed to a National Wildlife Refuge. 



i< to 



1£ 18 



Kure Atoll designated a State Wildlife Sanctuary. 

State of Hawaii Department of Land and Natural 
Resources was assigned management 
responsibilities for Kure Atoll. 



1! 33 



Northwestern Hawaiian Islands Coral Reef Ecosystem 
Reserve established by President Bill Clinton. 

NOAA's National Ocean Service was assigned 
stewardship responsibilities for the NWHI Coral Reef 
Ecosystem Reserve. 



Papahanaumokuakea Marine National Monument 
established by President George W. Bush. 

Department of Commerce through NOAA, Department 
of Interior through USFWS, and State of Hawaii 
through the Department of Land and Natural 
Resources were assigned Co-Trustee and day-to-day 
responsibilities for managing all land and marine 
areas of the Northwestern Hawaiian Islands. 



Hawaiian Islands Bird Reservation established by 
President Theodore Roosevelt. 

U.S. Department of Agriculture was assigned 

stewardship responsibilities for the islets and reefs 

of the Northwestern Hawaiian Islands. 



Midway Atoll National Wildlife Refuge established by 

President Ronald Reagan. 

U.S. Fish and Wildlife Service was assigned 
stewardship responsibilities for Midway Atoll. 



1£ 16 



Midway Atoll administration transferred to U.S. Fish 
and Wildlife Service by President Bill Clinton. 

USFWS assumed full management responsibilities for 
Midway Atoll after 1993 decommission of Midway 

Naval Air Facility. 



2( )0 



r 



2C )5 



Northwestern Hawaiian Islands State Marine Refuge 
established by Governor Linda Lingle. 

State of Hawaii Department of Land and Natural 

Resources was assigned stewardship 

responsibilities for near-shore marine areas of the 

Northwestern Hawaiian Islands. 



2( )6 



Figure 10.13. History of Management in the NWHI. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

MONUMENT PERMIT APPLICATIONS AND PERMIT ISSUANCE 

Since the designation of the Monument, all activities conducted within the Monument boundaries must meet 
the findings of the Proclamation and obtain a permit from the Monument. The Monument permitting program is 
one of the management tools that the Co-Trustees use to regulate the potential impacts of human activities on 
the Monument resources. Prior to the establishment of the Monument, the separate agencies responsible for 
management of the NWHI had separate permit applications, reviews and issuance processes and their own 
permit reporting requirements. Under Co-Trustee management, activities in the NWHI are prohibited, with lim- 
ited exception, unless authorized by a Monument permit. Applications for all applicable activities are reviewed 
and permits issued jointly by the three Co-Trustee agencies. 

A joint Monument permit application template and review process were developed and implemented in 2007. 
All applications are reviewed by managers, scientists, other experts within the three Monument Co-Trustee 
agencies and by Native Hawaiian cultural reviewers. In addition, summaries of permit applications are posted 
for public notification, and all applications for activities in State waters must be posted in full for public review 
before they are considered for approval by the State of Hawaii Board of Land and Natural Resources Land 
Board. 

In order for a project to be permitted, it must comply with National Environmental Policy Act requirements 
and all other federal and state required permits and consultations. In addition, any permitted activity must 
meet all of the Findings of the Presidential Proclamation (Proclamation 8031) establishing the Monument. 
Information on Monument permit application procedures is available at http://papahanaumokuakea.gov/re- 
source/permits.html. 

In addition to meeting the findings in the Proclamation, proposed activities to be conducted in the Midway 
Atoll SMA and the other National Wildlife Refuge areas, proposed activities continue to be subject to findings 
of appropriateness (603 FW 1) and compatibility determinations (16 U.S.C. 668dd-668ee and 603 FW 2) by 
USFWS to ensure the activities meet the purposes for establishing the Hawaiian Islands and Midway Atoll 
National Wildlife Refuges and the mission of the National Wildlife Refuge System. 



Findings of Presidential Proclamation 8031 

□ The activity can be conducted with adequate safeguards for the resources and ecological integrity of the Monu- 
ment. 

□ The activity will be conducted in a manner compatible with the management direction of the Proclamation, consid- 
ering the extent to which the conduct of the activity may diminish or enhance Monument resources, qualities, and 
ecological integrity; any indirect, secondary, or cumulative effects of the activity; and the duration of such effects. 

□ There is no practicable alternative to conducting the activity within the Monument. 

□ The end value of the activity outweighs its adverse impacts on Monument resources, qualities, and ecological in- 
tegrity. 

□ The duration of the activity is no longer than necessary to achieve its stated purpose. 

□ The applicant is qualified to conduct and complete the activity and mitigate any potential impacts resulting from its 
conduct. 

□ The applicant has adequate financial resources available to conduct and complete the proposed activity and miti- 
gate any potential impacts resulting from its conduct. 

□ The methods and procedures proposed by the applicant are appropriate to achieve the proposed activity's goals in 
relation to their impacts to Monument resources, qualities, and ecological integrity. 

□ The applicant's vessel has been outfitted with a mobile transceiver unit approved by NOAA Office of Law Enforce- 
ment and complies with the requirements of Proclamation 8031. 

□ There are no other factors that would make the issuance of a permit for the activity inappropriate. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Types of Permits Issued 

Applicants can apply for Monument permits under the following six permit categories: Research, Conservation 

and Management, Education, Native Hawaiian Practices, Recreation and Special Ocean Use. 

Research 

Research permits are authorized for those activities that enhance the understanding of Monument resources 
and improve resource management decision making. Priority is given to research proposals that help to meet 
the management needs of the Monument Co-Trustee agencies. Examples of types of activities issued under 
a research permit include biological inventories, ecosystem-based research, benthic mapping, habitat charac- 
terization, restoration investigations, cultural studies, and terrestrial and marine archaeological research. 

Conservation and Management 

Conservation and Management permits are authorized for those activities that are required for general man- 
agement of the Monument. This may include activities associated with resource management, such as field 
station operations, marine debris removal, development and maintenance of infrastructure, species and habitat 
restoration, and long-term resource monitoring programs such as monitoring of endangered species, seabird 
populations, and terrestrial native plant communities. Conservation and Management permits also provide a 
mechanism enabling rapid response and follow-up to critical events in the Monument that cannot be antici- 
pated, such as vessel groundings, coral bleaching episodes and invasive species detection. 

Education 

Education permits are authorized for those activities that further the educational value of the Monument. These 
activities may enhance the understanding of ecosystems, improve resource management decision making, 
promote Native Hawaiian knowledge and values, or aid in enforcement and compliance efforts. Priority is given 
to those activities that have clear educational or public outreach benefits and that promote "bringing the place 
to the people, rather than the people to the place." Examples of past projects issued under an education permit 
include teacher-at-sea programs, distance learning projects, and university classes. 

Native Hawaiian Practices 

Activities conducted under a Native Hawaiian Practice permit must be noncommercial, deemed appropriate 
and necessary by traditional standards, benefit the NWHI and Native Hawaiian community, perpetuate tradi- 
tional knowledge and restrict the consumption of harvested resources from the Monument. Examples of activi- 
ties permitted under a Native Hawaiian Practice permit include the entry of vessels for the purpose of applying 
and transferring knowledge of traditional navigation techniques and conducting ceremonies at historic cultural 
sites on Nihoa or Mokumanamana. Permit conditions and protocols for Native Hawaiian Practice permits 
will continue to be developed by the Monument Management Board, including the Office of Hawaiian Affairs 
through consultation with the Native Hawaiian Cultural Working Group and the Native Hawaiian community. 

Recreation 

Recreation permits are limited to the Midway Atoll Special Management Area in the Monument. Recreational 
activities may not be associated with any for-hire operation or involve any extractive use. Examples of recre- 
ational activities that may be permitted include snorkeling, SCUBA diving, wildlife viewing and kayaking. 

Special Ocean Use 

Special Ocean Use permits are authorized for those projects related to commercial ocean uses, such as eco- 
tourism or documentary filmmaking, that have a demonstrated net benefit to the Monument. Special Ocean 
Use is defined as any activity or use of the Monument that will generate revenue or profits for one or more of 
the persons associated with the activity or use. Activities that could potentially qualify as another permit type 
but that directly generates revenue or profit for at least one of the persons involved in the activity can only 
be permitted as Special Ocean Use. In addition, Special Ocean Use proposals involving activities outside of 
the Midway Atoll Special Management Area must have demonstrated educational or research purposes that 
directly benefit the conservation and management of the Monument. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

Emergencies, Law Enforcement Activities and Armed Forces Actions 

Permits are not required for those activities conducted within the Monument that are necessary to respond 
to emergencies or that are necessary for law enforcement purposes. Activities and exercises of the Armed 
Forces (including those carried out by the USCG) do not require a permit but must be conducted consistent 
with applicable federal laws. All other human presence, including activities conducted by the Co-Trustee agen- 
cies, require review and approval through the Monument permitting process. 

Transit Without Interruption Through the Monument 

Uninterrupted passage by vessels through the Monument does not require a permit but vessel operators must 
provide official notice prior to entering and upon departing. Official notification ensures that managers know 
at any given time who is present in the Monument either conducting activities under an authorized permit or 
transiting without interruption 

Additional Federal and State Permits and Consultations Required for Work in the Monument 
In addition to the permit requirements of the Monument, several other federal and state permits and/or consul- 
tations are required for many of the activities conducted in the NWHI. For example, all personnel working with 
threatened or endangered species must obtain an endangered species permit. Anyone handling any bird spe- 
cies must obtain one or more permits from the U.S. Fish and Wildlife Service Office of Migratory Bird Manage- 
ment, and all scientists working with marine mammals must obtain one or more permits from the NOAA Fish- 
eries Office of Protected Resources. Consultations may also be required as described under the U.S. ESAor 
Environmental Protection Agency regulations. Finally, although bottomfishing within the Monument boundaries 
will be phased out in 2011, all current bottomfishing operations are required to have valid federal fishing per- 
mits and state commercial marine licenses and fishing vessel registrations to operate within the Monument 



Table 10.7. Numbers of Monument permits granted, by permit type, for ac- 
tivities conducted in 2007. Numbers of projects that were newly-initiated in 
2007 and renewal projects (ongoing or long-term projects initiated in previ- 
ous years) are also listed. 



2007 Permitted Activities Conducted Within the Monument 
The first full year in which permits were 
issued by the Monument was 2007. 
Prior to June 2007, the State of Hawaii 
issued separate State permits for the 
Monument. Of a total of 51 permitted 
projects in the Monument in 2007, six 
were issued both Monument and state 
permits. The remaining 45 projects 
were issued a single joint Monument 
permit, issued by all three Co-Trustee 
agencies. Table 10.7 presents informa- 
tion on the number of permits issued, 
by permit type, for activities conducted 
in the Monument in 2007. The numbers 
of newly permitted projects and renewal 
projects (i.e., ongoing or long-term proj- 
ects initiated in previous years) are shown. 

Human activity in the NWHI has been greatly reduced relative to the height of military activity associated with 
World War II. Between 1940 -1945 more than 3,000 people were stationed at Midway Atoll and approximately 
125 were stationed at Tern Island wintin French Frigate Shoals. (Amerson, 1971). The overall level of human 
presence in the Monument in 2007 is indicated in Table 10.8. Eighteen ship cruises and a total of 99 flights 
transported permitted personnel and supplies to and from the Monument. The average number of people 
aboard ship per day throughout the year was 32, and the average number of people on land per day through- 
out the Monument was 83, for a total average of 115 people in the Monument per day in 2007. The number of 
people on land per day was highest at Midway Atoll, with an average human population of 70. Human presence 
on all other islands and atolls was an order of magnitude lower, with an average of 6.3, 4.2, and 1.5 people on 
land per day at French Frigate Shoals, Laysan Island and Kure Atoll, respectively, and on average, less than 



PERMIT TYPE 


NUMBER OF 
MONUMENT PER- 
MITS GRANTED 


NUMBER OF 
NEW NWHI 
PROJECTS 


NUMBER OF 

RENEWAL NWHI 

PROJECTS 


Research 


37 


16 


21 


Conservation and 
Management 


5 





5 


Education 


2 


2 





Native Hawaiian 
Practices 


1 


1 





Recreation 


1 


1 





Special Ocean Use 


5 


5 





TOTAL 


51 


25 


26 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



one person per day on all other islands 
and atolls in the chain. 

The following map (Figure 10.14) indi- 
cates locations where permitted activi- 
ties occurred in the Monument in 2007. 
Many of the permits issued allowed for 
work to be conducted at multiple loca- 
tions. Thus, for example, a single permit 
may have included work only at French 
Frigate Shoals, or it may have allowed 
for visits to all islands and atolls. In 2007 
the majority of the activities occurred at 
Midway Atoll, Pearl and Hermes Atoll 
and French Frigate Shoals. Midway 
Atoll and French Frigate Shoals have 
relatively easy access and the infra- 
structure to support activities, includ- 
ing landing strips and facilities to house 
year-round personnel. 

In order to assess cumulative impacts, it 
is important for managers to understand 
the trends and patterns of the different 
types of permitted activities that have 
occurred in the Monument. The follow- 
ing section spatially represents the dis- 
tribution of activities that took place in 
the Monument by the types of permits 
issued in 2007 



Table 10.8. Number of ship cruises and flights, and average number of 
people on land per day in the Monument in 2007. 



TRANSPORTATION 


Number of Ship Cruises 


18 


Number of Flights 




French Frigate Shoals 


13 


Midway Atoll 


86 


VISITATION 






Average Number of People on Land per Day 


83 


Nihoa 


0.02 


Mokumanamana 


0.06 


French Frigate Shoals 


6.30 


Laysan 


4.20 


Lisianski 


0.30 


Pearl and Hermes Atoll 


0.80 


Midway Atoll 


70.0 


Kure Atoll 


1.50 


Average Number of People on Ships per Day 


32 


Average Number of People in Monument per Day 


115 




\ J Papahanaumokuakea 

Marine National Monumenl Boundary 



100 200 300 400 

Kilonraters 



Figure 10.14. Locations of all permitted activities within the Monument in 
2007. Map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Research Activities by Location 
Islands and atolls with the highest levels 
of permitted Research activities in 2007 
included French Frigate Shoals, Pearl 
and Hermes Atoll, and Midway Atoll (Fig- 
ure 10.15). Non-emergent banks and 
reefs, including Twin Banks, St. Roga- 
tien and Brooks Banks, and Maro Reef, 
saw the lowest levels of research activi- 
ties, while Mokumanamana and Gard- 
ner Pinnacles had the fewest number of 
Research activities conducted on emer- 
gent lands. Managers can use informa- 
tion on the distribution of past research 
activities to better plan and target future 
research and to ensure that data gaps 
are filled for those areas for which with 
less information is available. 

Conservation and Management 
Activities by Location 
In 2007, U.S. Fish and Wildlife Service 
Conservation and Management activi- 
ties took place at Nihoa, French Frigate 
Shoals, Laysan, and Pearl and Hermes 
Atoll (within Hawaiian Islands National 
Wildlife Refuge), and at Midway Atoll 
(Midway Atoll National Wildlife Refuge). 
State of Hawaii DLNR activities took 
place at Kure Atoll, and NOAA PIFSC- 
CRED marine debris removal activities 
occurred at French Frigate Shoals, Lay- 
san, Lisianski, Pearl and Hermes Atoll, 
and Kure Atoll (Figure 10.16). 




Figure 10.15. Locations of permitted Research activities in 2007. Several 
of the 37 Research permits authorized work at multiple locations within the 
Monument; thus, the total number of permits in the figure below adds to 
more than 37. Map: K. Keller. 




i _J Papahanaumakuakea 

Marine National Monumenl Boundary 



100 200 300 400 

D KilO ill C10"i 



Figure 10.16. Locations of permitted Conservation and Management activi- 
ties in 2007. Two of the five Conservation and Management permits autho- 
rized work at multiple locations within the Monument; thus, the total number 
of permits in the figure below adds to more than five. Map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Education Activities by Location 
NOAA Monument education activities in 
2007 took place almost entirely aboard 
ship and during 12 shallow-water free 
dives, with land visits made only at Mid- 
way Atoll and Kure Atoll. Sites visited, 
where photos and video footage were 
taken, included Nihoa, Gardner Pin- 
nacles, Laysan, Lisianski, Pearl and 
Hermes Atoll, Midway Atoll and Kure 
Atoll. 



DLNR education activities took place 
aboard ship and during 13 shallow- 
water dives at French Frigate Shoals, 
Pearl and Hermes Atoll, and Midway 
Atoll (Figure 10.17). Photos and video 
footage were taken at each of these 
sites. 




■ _J Papahanaumci-kuSkea 

Marine National Monument Boundary 
N 
100 200 300 400 

3 Kilometers 



Figure 10.17. Locations of permitted Education activities in 2007. The two 
Education permits authorized work at multiple locations within the Monu- 
Native Hawaiian Practices Activities by ment; thus, the total number of permits in the figure below adds to more 
Location tnan two - Map: K. Keller. 

A single Native Hawaiian Practices per- 
mit was issued in 2007, to the University 
of Hawaii. Activities conducted under 
this permit included an overnight stay on 
Mokumanamana to conduct traditional 
ceremonies in observance of the sum- 
mer solstice, and a stop off of the island 
of Nihoa to conduct additional ceremo- 
nies aboard ship (Figure 10.18). 




Legend 



Number of 2007 Native Hawaiian Practices 
Permit Activities 
• 1 

I Papahanaurnokuakea 
Marine National Monument Boundary 
N 
100 ZOO 300 100 

Kilometers 



Figure 10.18. Locations of permitted native Hawaiian practices and activi- 
ties in 2007. Map: K. Keller. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



Special Ocean Use Activities by Location 
Five Special Ocean Use permits were 
issued in 2007 occurring at Midway and 
French Frigate Shoals (Figure 10.19). 
Two of these permits were associated 
with the commemoration of the 65th An- 
niversary of the Battle of Midway and 
involved flight and ship transportation of 
World War II veterans and their families 
to Midway Atoll for the one-day celebra- 
tion. A third Special Ocean Use permit 
was issued to the British Broadcasting 
Corporation for high-definition filming of 
tiger shark predation on albatross fledg- 
lings at French Frigate Shoals. The final 
two Special Ocean Use permits were 
issued for two individuals to conduct 
filming and still photography associated 
with NOAA Education and Research ac- 
tivities. 




Number of 2007 Special Ocean Use 
Permit Activities 
• 2 



' ] Papahanaumokuakea 

Marine National Monument Boundary 
N 

a toe 200 300 400 

Kilometers 



Figure 10.19. Locations of permitted Special Ocean Use activities in 2007. 
Map: K. Keller. 



Table 10.9. Relative size and visitation rates for Hanauma Bay, Yosemite 
National Park, Great Barrier Reef Marine Park and Papahanaumokuakea 
Marine National Monument. 



PROTECTED AREA PER DAY SIZE 



Hanauma Bay Nature Preserve 100 acres 



Yosemite National Park 



760,000 
acres 



Great Barrier Reef Marine Park 



85,100,000 
acres 



Papahanaumokuakea Marine 
National Monument 



89,500,000 
acres 



VISITATION PEOPLE 

RATE PER ACRE 



3,000 people/day 



30 



9,600 people/day 



0.01 



PMNM Relative Size and Visitation Rates 
Papahanaumokuakea Marine National 
Monument is the largest protected area 
in the United States, and is larger than 
all of the U.S. National Parks combined. 
It is also larger than almost all other 
protected marine areas in the world, in- 
cluding Great Barrier Reef Marine Park. 
Because of the protections put in place 
by Presidential Proclamation 8031, fol- 
lowed by the permitting system estab- 
lished by the Monument Management 
Board in 2007, the visitation rate to 
the Monument was significantly lower 
in 2007 compared to visitation rates to 
Hanauma Bay Nature Preserve, Yosem- 
ite National Park, and even Great Barrier Reef Marine Park (Table 10.9). The following figure (Figure 10.20) 
also illustrates the scale of Papahanaumokuakea Marine National Monument relative to western states on the 
mainland U.S. The Monument is larger than 46 of the 50 states, and if overlain on the west coast would span 
an area from Arizona through Nevada, Oregon, and Washington state. 



5,423 people/day 



115 people/day* 



0.00006 



0.000001 



Includes 70 people per day at Midway Atoll. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



N 



Olympic National Park l^ 








Kure Atoll SPA 1 








Midway Island SMA 1 



Papahanaumokuakea Marine National 
Monument and US West Coast 



s **,, 



r *»« : 




Yosemite National Park 



Miles 
50 100 200 
I 1— I 1 1 1 1 1 1 



Papahanaumokuakea Marine National Monument Boundary 
Special Protection/Management Area 
National Park 



Nihoa Island SPA 



W^cson / 



Figure 10.20. Size of the Monument relative to states on the west coast of the mainland U.S. Source: D. Turner, PMNM. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

FUTURE DIRECTIONS AND IMPLICATIONS FOR A BIOGEOGRAPHIC ASSESSMENT TO 
SUPPORT HAWAIIAN ARCHIPELAGO SPATIAL MANAGEMENT 

A primary use of implementing a marine biogeographic assessment for the NWHI was to further the under- 
standing of the temporal and spatial coupling between the oceanographic characteristics, pelagic and benthic 
habitats, living marine resources and human uses that collectively comprise the NWHI ecosystem. The biogeo- 
graphic assessment process is defined in this document's introduction and illustrated in Figure 10.21. Much 
of the data and information synthesized for this investigation addressed an area (e.g., oceanographic charac- 
teristics) much greater than the boundaries of the Monument or the individual atolls that comprise the island 
chain. However, the study area did not encompass the entire Hawaiian Archipelago that would place in context 
the biogeographic results for the NWHI when compared to the Main Hawaiian Islands (MHI). The exception is 
the work that has been done to characterize the shallow-water reef fish assemblages around both the heavily 
fished MHI and the very limited exploited reef fishes of the NWHI. 




Figure 10.21. Generalized biogeographic assessment process developed by CCMA-BB. Source: Kendall and Monaco, 
2003. 

The 2,500 km long Hawaiian Archipelago is unified by its geologic origin and geographic isolation. This vast 
area is subject to great spatial gradients in oceanography, erosion and geomorphology. The Hawaiian ma- 
rine ecosystem has some of the highest marine endemism on the planet, with many species unique to the 
archipelago. Given these broad-scale characteristics, the recently published NOAA Technical Memorandum 
(NOAA, 2008), "Hawaiian Archipelago Marine Ecosystem Research" plan (HAMER), noted that Hawaii could 




w 



NOAA Tetfinicai .Memorandum NMFS-PiFSC-w 
February 2008 



Hawaiian Archipelago Marine Ecosystem Research 
(HAMHR) 







1 1 . ."■■- . . liivisiun of Aquatic EuCSOUfCei 

Papoh9naumnki4kca Marim; National Monument 

NOAA Pacitiu Islands l-'i^hcnw Sticncfl Cento 

University of Hawaii 

U.S. Fish and Wildlife Service 

Wtotmi Pacific Hcijional Fishery ManamsmcrH Council 



Pacific islands Fusnwios Science Center 

National Marine F*h*ne» Sann 

National Oceanic find Atmospheric Ai!>wirsirit:>dri 
U.S OepanjTOrvi of Commeree 



A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 

serve as "a large scale archipelagic laboratory for the investigation of bio- 
physical processes, comparing the protected and nearly pristine NWHI to 
the heavily used MHI to improve resources management in Hawaii and in 
comparable marine ecosystems worldwide (NOAA, 2008; Figure 10.22). 
HAMER identified six research themes important to the management of the 
Hawaiian Archipelago. They are: 

Ecosystem Indicators and Metrics; 
Native Biodiversity and Invasive Species; 
Connectivity; 
Human Interactions; 
Resilience and Recovery; and 
Modeling and Forecasting. 

Consistent with HAMER, the draft Papahanaumokuakea Marine National 
Monument Natural Resources Science Plan (NRSP), specifically cited a 
need for archipelagic-wide homogeneity in research planning and execu- 
tion and adopted nearly identical thematic focus areas: 

Ecological Processes and Connectivity; 

Biodiversity and Habitats; 

Human Impacts; 

Indicators of Change and Monitoring; and 

Models and Forecasting. 

Although HAMER is a plan for implementation over a ten year period, and the NRSP spans a shorter, five 
year period. Both plans provide the spatial and institutional framework to conduct research and integrate data 
and information in support of management of the Hawaiian Archipelago as a single, interconnected entity. 
With the recent increase in ecosystem monitoring data (e.g., Hawaii Coral Reef Assessment and Monitoring 
Program [CRAMP], MHI and NWHI Reef Assessment and Monitoring Programs [MHI/NWHI RAMP]), benthic 
habitat mapping (NOAA, 2007), and the assessment of habitats and associated fishes in and outside of marine 
protected areas (Friedlander et al., 2005, 2007), our ability to conduct archipelagic-wide research syntheses 
and biogeographic assessments is greatly enhanced. The biogeographic assessment process and this NWHI 
product directly support these multi-agency visions of HAMER and the PMNM NSRP, and provide a key start- 
ing point and many spatial data products that could be incorporated in a recommended MHI or archipelago- 
wide biogeographic assessment. 



Figure 10.22. The Hawaiian Ar- 
chipelago Marine Ecosystem Re- 
search. 




A Marine Biogeographic Assessment of the Northwestern Hawaiian Islands 



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